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ON  THE  DEVELOPMENT,  MORPHOLOGY, 

AND  ECONOMIC  IMPORTANCE 

OF  CHICKEN  CESTODES 

Morphology  of  Adult  and  Larval  Cestodes  from  Poultry 

Studies  on  the  Transmission  and  Prevention  of  Cestode 
Infection  in  Chickens 

BY 
JOHN  EARL  GUTBERLET 

A.  B.  Bethany  College,  Kansas,  1909 
A.  m;  University  of  Illinois,  1911 


THESIS 


lubmitted  in  Partial  Fulfillment  of  the  Requirements  for  the 

Degree  of 
DOCTOR  OF  PHILOSOPHY 


IN 

THE   GRADUATE   SCHOOL 
OF  THE 

UNIVERSITY  OF  ILLINOIS 

1914 


ON  THE  DEVELOPMENT,  MORPHOLOGY, 

AND  ECONOMIC  IMPORTANCE 

OF  CHICKEN  CESTODES 

Morphology  of  Adult  and  Larval  Cestodes  from  Poultry 

Studies  on  the  Transmission  and  Prevention  of  Cestode 
Infection  in  Chickens 

BY 
JOHN    EARL   GUTBERLET 

A.  B.  Bethany  College,  Kansas,  1909 
A.M.  University  of  Illinois,  1911 


THESIS 


Submitted  in  Partial  Fulfillment  of  the  Requirements  for  the 

Degree  of 
DOCTOR  OF  PHILOSOPHY 

'' .  •  ,      IN   ■  " 

m 

THE   GRADUATE   SCHOOL 
OF  THE 

UNIVERSITY   OF   ILLINOIS 

1914 


24  john  e.  gutberlet 

Structure  of  Adult  and  Larva  (Cysticercus) 

A.  adult 

Choanotcenia  infundibuliformis  (Goeze  1782)  Railliet  1896 

1.  Diagnosis:  Length  50  to  200  mm.  Scolex  (Fig.  2)  small, 
rounded,  or  conoidal,  about  0.4  mm.  wide.  Rostellum  (Fig.  2,  3,  r) 
60  to  70ix  in  diameter,  armed  with  a  single  row  of  16  to  20  hooks 
(Fig.  8)  25  to  30ju,  long,  with  long  dorsal  root  and  short  ventral 
root.  Suckers  prominent,  elongated  antero-posteriorly,  length  180 
to  210/x;  breadth  135  to  175/x  between  the  extreme  outer  edges. 
Neck  short  and  unsegmented,  somewhat  narrower  than  broad.  In 
specimens  well  extended  neck  much  narrower  than  head.  Anterior 
proglottids  very  short  and  as  they  become  older  funnel-shaped, 
much  narrower  at  anterior  than  at  posterior  margins;  posterior 
segments  1.5  to  2.5  mm.  broad  and  1.5  to  3  mm.  long  according  to 
amount  of  contraction,  with  convex  lateral  borders,  nearly  as  wide 
at  anterior  as  at  posterior  margin.  Genital  pores  irregularly  alter- 
nating, situated  one  in  each  segment  in  the  anterior  third  of  the 
lateral  margin,  usually  under  cover  of  the  backward  projecting  bor- 
der of  the  preceding  segment.  Vas  deferens  (Fig.  14,  vd)  and 
vagina  pass  between  excretory  canals  and  dorsal  to  nerve  trunk. 

Male  Reproductive  Organs:  Testicles  (Fig.  14,  t)  25  to  40 
or  more,  60  in  some  cases,  in  posterior  half  of  proglottid,  posterior 
and  lateral  to  large  yolk  gland,  within  limits  of  excretory  canals. 
Vas  deferens  passes  forward  and  in  anterior  third  of  proglottid 
forms  a  mass  of  coils  between  ovary  and  excretory  vessels  from 
which  it  extends  outward  as  a  convoluted  tube  to  base  of  cirrus 
pouch.  Cirrus  pouch  (Fig.  14,  15,  cp)  ovoid  in  shape,  75  to  95fi 
in  long  diameter.  Portion  of  vas  deferens  in  cirrus  pouch  is  much 
coiled.  Cirrus  50  to  65ix  long,  armed  with  spines ;  outer  surface  of 
cirrus  pouch  forms  base  of  deep  genital  cloaca. 

Female  Reproductive  Organs:  Vaginal  opening  in  genital 
cloaca  posterior  to  cirrus.  Vagina  posterior  to  cirrus  pouch,  after 
crossing  ventral  excretory  canal  dilated  to  form  ovoid  seminal 
receptacle,  posterior  and  ventral  to  vas  deferens,  extending  to  well 
developed  shell  gland,  40  to  50/i,  in  diameter  located  in  front  of 
middle  of  proglottid.     Transversely  elongated  ovary    (Fig.   14,  o) 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  25 

occupies  anterior  portion  of  middle  field  of  proglottid  in  front  of 
shell  gland.  Large  yolk  gland  posterior  to  ovary  and  shell  gland, 
irregular  in  shape,  elongated  transversely,  with  convex  ventral  sur- 
face and  concave  dorsal  surface.  Uterus  (Fig.  16,  ti)  developed 
as  tube  between  anterior  and  ventral  lobes  of  ovary.  Gravid  uterus 
fills  up  most  of  proglottid,  extending  beyond  excretory  canals  on 
each  side.  Eggs  oval  (Fig.  7),  with  very  thin  membrane  next  em- 
bryo, followed  by  thick,  smooth  membrane  40  by  32[x  to  45  by  36/* 
in  diameter,  and  one  or  two  outer  membranes,  very  thin  and  wrink- 
led in  preserved  material.  Diameter  of  outer  membrane  65  by  40/* 
to  60  by  45ft ;  at  each  pole  of  outer  membrane  a  delicate  appendage. 
Embryonal  hooks  18/*  long.     Embryo  32  by  22ju,  in  diameter. 

2.  Morphology:  The  scolex  of  the  living  worm  shows  up 
very  prominently  and  can  be  used  as  a  distinguishing  feature. 
When  first  removed  from  the  intestinal  wall  the  suckers  appear 
distinct  and  the  neck  is  much  narrower  than  the  scolex.  Soon 
after  the  removal  it  often  contracts  and  takes  on  the  appearance  of 
a  flattened  bulb  which  includes  the  neck  and  anterior  segments 
(Fig.  1).  This  feature  is  characteristic  of  this  species  and  is  a 
factor  which  alone  assists  very  materially  in  distinguishing  it  from 
others  that  occur  in  chickens. 

The  rostrum  or  crown  of  the  scolex  is  somewhat  pointed  when 
the  rostellum  is  enclosed  within  its  sheath  (Fig.  2).  The  rostellum 
is  an  ovoid  structure  with  a  bulbous  expansion  at  its  anterior  end. 
It  has  a  length  of  140|U.  and  a  breadth  of  60  to  65/i  at  its  anterior 
end.  A  crown  of  18  hooks  is  arranged  in  a  single  row  around  the 
bulbular  anterior  end.  The  structure  of  the  wall  is  of  a  fibrous 
nature  and  presents  a  transversely  striated  appearance  due  to  con- 
traction. In  the  interior  of  the  rostellum  the  structure  is  a  con- 
nective tissue  mass  with  few  cells,  some  of  which  possess  long 
processes.  The  hooks  (Fig.  8)  are  SO/n  in  length  with  a  long  dorsal 
root  and  a  short  ventral  root. 

The  rostellar  sheath  or  sac  (Fig.  3,  rs)  into  which  the  rostel- 
lum is  withdrawn  is  oval  in  shape  and  230  to  240/a  in  length  by  80 
to  90)u,  in  width  at  its  broadest  point.  Histologically,  the  structure 
is  that  of  a  fibrous  connective  tissue  type  with  spherical  and  spindle- 
shaped  cells.     The  cells  coming  in  contact  with  the  rostellum.  as 


26  JOHN    E.    GUTBERLET 

well  as  those  on  the  outer  edge  of  the  sac,  bear  long  processes.  The 
outer  layer  of  the  rostellar  sac  is  composed  of  longitudinal  and 
oblique  fibers  of  a  muscular  nature  which  probably  have  for  their 
function  the  movement  of  the  rostellum. 

The  four  excretory  canals,  that  have  extended  forward  through 
the  entire  length  of  the  body,  unite  in  the  scolex  to  form  a  ring 
(Fig.  3,  e.v),  which  lies  in  the  tissue  of  the  rostellar  sac  around  the 
body  of  the  rostellum. 

The  suckers  are  prominent.  They  are  oval  in  shape  and  in 
preserved  specimens  measure  180  to  210/^  in  length  and  from  135 
to  175jM  in  extreme  breadth.  In  the  center  of  each  sucker  there  is 
a  depression  or  an  acetabulum,  30  to  40/^  in  diameter.  The  entire 
inner  surface  of  the  suckers  possesses  minute  booklets  or  spines 
(Fig.  4)  1.5  to  2(1  long.  These  booklets  not  only  line  the  suckers 
but  also  extend  over  the  entire  surface  of  the  scolex  (Figs.  3,  5) 
and  down  onto  the  neck  region ;  they  disappear  before  reaching 
the  first  segment.  They  appear  more  distinctly  on  scolices  that  are 
somewhat  contracted  than  on  those  that  are  well  extended.  These 
booklets  can  be  seen  only  in  sections  as  they  are  too  small  to  be 
distinguished  readily  in  whole  mounts. 

Musculature:  The  longitudinal  muscle  fibers  are  arranged  in 
bundles  which  are  scattered,  forming  a  loose  irregular  layer.  The 
bundles  are  numerous  and  nearly  of  a  uniform  size.  There  are  no 
transverse  muscle  fibers  present  except  a  few  minute  oblique  fibers 
which  connect  some  of  the  longitudinal  fibers  near  the  ends  of  the 
proglottids.  Some  dorso-ventral  fibers  are  present,  but  they  are 
not  abundant. 

Nervous  System:  The  longitudinal  nerve  fibers  are  arranged 
in  fiber  tracts  which  approach  the  structure  of  a  nerve  cord.  The 
individual  fibers  do  not  form  a  compact  mass,  but  are  more  or  less 
free  in  the  tract.  Nerve  cells  have  no  definite  arrangement,  but 
are  situated  irregularly  along  the  fiber  tract  (Fig.  6).  The  nerve 
cells  are  somewhat  spindle-shaped  and  quite  large,  being  from  20 
to  25,a  long  by  6  to  8/x  wide  with  large  nuclei.  Transverse  nerves 
are  composed  of  individual  cells  with  long  processes  extending 
transversely  from  the  lateral  fiber  tracts.  The  transverse  fibers  are 
much  scattered  and  have  no  definite  arrangement  except  that  they 


MORrHOLOGY   OF    CESTODES    FROM    POULTRY  27 

are  more  numerous  near  the  ends  of  the  proglottids.  Peripheral 
nerve  cells  are  widely  and  irregularly  distributed.  They  are  more 
numerous  at  the  anterior  end  of  the  proglottids,  especially  on  the 
portion  that  is  covered  by  the  backward  extension  of  the  preceding 
segment. 

Excretory  System:  The  excretory  system  is  fairly  well  devel- 
oped in  this  form.  The  ventral  canal  (Fig.  14,  v  ex)  is  the  larger, 
and  has  a  diameter  of  28  to  30/x.  A  transverse  canal  unites  the 
two  longitudinal  canals  in  each  segment.  The  dorsal  canals  (Fig. 
14,  d  ex)  are  much  smaller,  having  a  diameter  of  6  to  8ju,  and  are 
not  united  by  transverse  connections.  The  four  longitudinal  canals 
extend  anteriorly  to  the  scolex  where  they  unite  to  form  a  ring 
which  lies  in  the  rostellar  sheath  around  the  body  of  the  rostellum. 
The  vas  deferens  and  vagina  pass  between  the  dorsal  and  ventral 
excretory  canals. 

Male  Reproductive  Organs:  The  testes  vary  in  number,  us- 
ually from  25  to  40,  but  in  a  few  cases  the  number  is  much  greater, 
being  as  high  as  55  or  60.  The  testes  are  quite  large,  being  from 
40  to  SSfi  in  diameter,  and  are  located  in  the  posterior  half  of  the 
proglottid  (Fig.  14,  t),  posterior  and  lateral  to  the  yolk  gland.  The 
testes  are  not  arranged  in  layers,  but  are  grouped  in  a  more  or  less 
compact  mass  almost  entirely  within  the  limits  of  the  excretory 
canals.  The  vas  deferens  (Fig.  14,  vd)  in  the  anterior  third  of 
the  proglottid  forms  a  coiled  mass  at  the  side  of  the  ovary,  from 
whence  it  passes  laterad  to  the  cirrus  pouch  as  a  convoluted  tube. 
The  portion  of  the  vas  deferens  inside  the  cirrus  pouch  is  coiled, 
varying  in  extent  in  different  specimens  (Figs.  14,  15).  The  vas 
deferens  passes  into  the  cirrus.  There  is  no  seminal  vesicle  formed 
by  the  vas  deferens  in  the  cirrus  pouch  nor  are  there  any  accumu- 
lations of  sperm  cells.  The  cirrus  pouch  (Fig.  15)  is  ovoid  in 
shape  and  is  from  75  to  90/i  in  diameter.  The  wall  is  made  up  of 
layers  of  fibers  which  are  both  circular  and  oblique,  forming  a 
basket-like  network  which  incloses  the  cirrus  and  a  portion  of  the 
vas  deferens.  The  outer  wall  of  the  cirrus  pouch  forms  the  inner 
wall  of  the  deep  genital  cloaca.  The  cirrus  is  a  compact  structure 
from  50  to  65^  long  and  Hned  with  spines.  It  is  a  slightly  curved 
structure   passing   from   the   cirrus   pouch  and   curving  posteriorly 


28  JOHN    E.    GUTBERLET 

toward  the  vagina  which  is  directly  posterior  to  it.  The  cirrus  was 
not  observed  extending  from  the  genital  cloaca,  but  was  noted  in 
some  specimens  curving  toward  the  vagina,  though  not  passing  into 
it.  A  few  sperm  cells  were  present  in  the  vas  deferens,  also  in  the 
vagina  and  the  seminal  receptacle. 

Female  Reproductive  Organs:  The  large  ovary  (Fig.  14,  o) 
lies  in  the  anterior  third  of  the  proglottid  and  extends  transversely 
across  the  segment.  It  has  a  length  of  300/i.  and  a  breadth  of  about 
75  or  80/1,  at  its  broadest  point.  It  is  irregular  in  shape,  being  com- 
posed of  a  number  of  lobes.  The  end  which  is  nearest  the  genital 
pore  is  smaller  than  the  other,  allowing  room  for  the  mass  of  coils 
of  the  vas  deferens,  the  vagina,  and  the  seminal  receptacle.  The 
ovary  is  concave  on  the  dorsal  surface  and  convex  on  the  ventral. 
On  the  dorsal  surface  of  the  end  nearest  the  genital  pore  is  located 
the  seminal  receptacle  and  the  vagina.  The  ova  are  large  and  very 
distinctly  shown  in  the  ovary  (Fig.  16).  Posterior  to  the  ovary  is 
the  large  yolk  gland  (Fig.  14,  16,  y)  which  lies  about  the  middle  of 
the  proglottid.  It  is  irregularly  elongate  in  shape  and  extends 
transversely  across  the  segment,  having  a  length  of  from  120  to 
ISOfi  and  a  breadth  of  from  35  to  50/i,.  Immediately  in  front  of  and 
dorsal  to  the  yolk  gland  and  posterior  to  the  ovary  is  the  shell  gland 
(Fig.  14,  sg)  which  is  slightly  ovoid  in  shape,  40  to  50/x  in  diameter. 
A  small  duct,  the  vitelline  duct  (Fig.  16,  v),  passes  from  the  yolk 
gland  through  the  shell  gland  from  which  it  receives  a  duct.  The 
combined  ducts  after  passing  through  the  shell  gland  unite  with  the 
oviduct  (Fig.  16,  ov)  which  appears  as  a  curved  tube  leading  from 
the  ovary.  These  united  tubes  or  ducts  pass  anteriad  and  slightly 
ventrad  into  the  uterus  which  develops  as  a  blind  tube  in  the  region 
of  the  ventral  lobes  of  the  ovary.  This  blind  tube  (Fig.  16,  «) 
grows  in  size  and  extends  transversely  across  the  segment.  As  it 
becomes  larger  the  tube  forms  pockets  which  extend  anteriorly  and 
posteriorly  and  also  dorsally,  until  it  takes  up  the  entire  mass  of 
the  proglottid  between  the  excretory  canals.  In  gravid  segments 
it  even  extends  beyond  the  excretory  canals.  A  small  tube  or  duct, 
which  is  really  the  end  of  the  vagina,  connects  the  seminal  recep- 
tacle with  the  yolk-shell  gland  duct  and  oviduct.  This  tube  serves 
to  carry  the  sperm  to  the  eggs  in  the  oviduct  for  fertilization.     The 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  29 

seminal  receptacle  (Fig.  16,  sr)  is  a  dilation  of  the  vagina  into  an 
oval  shaped  structure  which  is  about  SO/x  long  and  from  25  to  30ju, 
in  breadth  at  the  widest  part.  From  the  seminal  receptacle  the 
vagina  passes  laterad,  lying  posterior  to  the  cirrus  pouch,  and  unites 
with  the  genital  cloaca.  The  genital  cloaca  has  its  pore  on  the 
lateral  margin  near  the  anterior  end  of  the  proglottid.  The  pore  is 
usually  covered  by  the  backward  projection  of  the  segment  anterior 
to  it.  The  vas  deferens  and  vagina  pass  between  the  dorsal  and 
ventral  excretory  canals  and  dorsal  to  the  nerve  tract.  The  vas 
deferens  is  dorsal  and  anterior  to  the  vagina. 

In  the  mature  segments  the  uterus  becomes  filled  with  ova  and 
it  increases  in  size  until  it  occupies  the  entire  area  between  the 
excretory  canals,  even  extending  beyond  the  canals  in  the  gravid 
proglottids.  The  uterus  finally  breaks  up  into  compartments,  each 
containing  a  single  embryo.  The  embryos  (Fig.  7)  are  about  32 
by  22/i,  in  diameter  with  onchospheric  hooks  18/x  long.  Usually 
three  membranes,  but  often  four,  enclose  the  embryo.  The  inner 
membrane  is  thin  and  closely  surrounds  the  embryo ;  the  next  is 
heavy,  being  from  1.5  to  2/i  thick,  composed  of  fibrous  layers  with 
a  few  cells  present.  This  layer  is  variable  in  thickness,  depending 
considerably  upon  the  amount  of  contraction  of  the  segment,  as  it 
ranges  in  size  from  40  to  32/x  to  50  by  26ix,  or  it  may  be  even 
slightly  larger.  Usually  one  (Fig.  7)  and  sometimes  two  thin  mem- 
branes are  found  on  the  outside  of  the  thick  layer.  These  are 
often  wrinkled  and  bear  at  each  end  an  appendage  formed  from 
the  outer  membrane  by  which  it  is  attached  to  the  wall  of  the  cap- 
sule or  compartment  of  the  uterus. 

In  this  species  the  oldest  proglottids  drop  off  from  the  worm 
before  they  are  fully  m.ature.  The  embryos  from  the  oldest  seg- 
ments on  the  worm  do  not  show  the  characteristics  of  entirely  ma- 
ture ones,  and  there  are  distinct  differences  between  them  and  those 
that  have  been  separated  from  the  worm  for  some  time.  Single 
proglottids  that  have  separated  from  the  worm  are  quite  active  and 
remain  in  the  intestine  for  some  time  before  passing  out  with  the 
feces.  Proof  of  this  is  furnished  by  the  fact  that  a  large  number 
of  the  free  proglottids  are  found  in  the  intestine  at  any  time.  Even 
tho  only  a  few  worms  are  present  in  the  intestine  of  a  bird  there  is 


30  JOHN    E.    GUTBERLET 

usually  a  large  number  of  free  proglottids.  If  they  did  not  remain 
in  the  intestine  for  a  considerable  length  of  time  there  would  not 
be  nearly  as  many.  Further  proof  is  furnished  by  the  fact  that  the 
free  proglottids  have  embryos  which  are  mature,  showing  the  oncho- 
spheric  characteristics,  while  the  oldest  segments  that  are  still  at- 
tached to  the  worm  have  embryos  that  are  not  entirely  mature. 
This  same  condition  has  been  observed  in  Davainca  proglottina  as 
Blanchard  (1891:435)  states  that  the  oldest  proglottids  separate 
from  the  others  and  remain  in  the  intestine  to  become  mature  be- 
fore passing  out.  The  proglottids  do  not  always  separate  from  the 
worm  singly,  but  may  drop  off  in  groups  of  three  or  four. 

The  fact  that  the  proglottids  separate  from  the  worm  before 
they  are  entirely  mature  is  one  of  great  importance  in  taking  up 
experimental  work  for  infection  of  intermediate  hosts.  If  the  em- 
bryos are  fed  to  insects  or  other  invertebrates  before  they  are  ma- 
ture they  will  be  digested,  and  thus  infection  cannot  be  produced. 

B.       CYSTICERCUS 

The  cysticercus  of  Choanotcenia  infundibiiliformis  was  found 
in  the  abdominal  region  of  the  body  cavity  in  the  common  house 
fly,  Mnsca  domestica.  The  flies  had  been  fed  on  embryos  from 
ripe  proglottids  of  this  species  of  worm,  and  at  the  end  of  twelve 
days  were  killed.  The  cysticerci  appear  to  be  nearly  ripe  or  ready 
for  transmission  into  the  adult  host.  The  time  for  the  develop- 
ment of  the  cysticercoid  varies  with  different  species  and  under 
different  conditions.  Grassi  and  Rovelli  (1892:85)  found  that 
Davainea  proglottina  developed  from  the  onchosphere  into  a  ripe 
cysticercus  in  less  than  twenty  days.  Schmidt  (1894:9)  found  that 
the  development  of  the  cysticercoid  of  Drepanidotcrnia  anatina 
(Krabbe)  varied  with  the  time  of  the  year  and  the  influence  of  the 
temperature.  In  the  summer  the  embryo  developed  in  an  ostracod, 
Cypris  ovata,  into  ripe  cysticercoids  in  two  weeks. 

The  cyst  proper  (Figs.  11,  12,  c)  containing  the  scolex  is  oval 
in  shape,  220/i,  long  and  120/x  in  diameter. 

The  bladder  (Fig.  12,  h)  ov  tail,  which  is  also  oval  in  shape, 
is  located  against  one  side  of  the  cyst  and  is  somewhat  flattened  on 
that  side.     It  is  220  to  2v30/x  long  and  from  116  to  120/t  in  breadth. 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  31 

The  scolex  is  SO/x  in  breadth  and  120jli  in  length;  neck  is  AOfx  in 
diameter  and  30  to  35/*  long;  suckers  are  55  to  60/x  in  diameter. 
The  rostellum  is  60/x  long  and  20/x  in  breadth,  armed  with  a  crown 
of  18  hooks  arranged  in  a  single  row.  These  hooks  (Fig.  9)  are 
30/*  long  with  a  long  dorsal  root  and  a  short  ventral  root.  The 
suckers  are  lined  with  numerous  minute  booklets  or  spines  1.5  to  2/* 
long  which  extend  over  the  edges  of  the  suckers  and  also  over  the 
greater  part  of  the  surface  of  the  scolex,  including  a  part  of  the 
neck  region.  Schmidt  (1894:  16)  described  cuticular  booklets  on 
the  suckers  of  Drepanidotmiia  anatina. 

The  size  of  the  scolex  may  be  somewhat  variable  as  shown  by 
those  in  the  cysticercoids  of  Drepanidotcunia  anatina  by  Schmidt 
(1894:  10).  In  that  species  the  intermediate  host  could  be  one  of 
two  or  more  species  of  crustaceans  and  the  size  of  the  cysticercoid 
varied  with  the  size  of  the  host  in  which  it  was  parasitic. 

The  head  of  the  rostellum  is  conical  in  shape,  bearing  a  bluntly 
pointed  apex  anterior  to  the  end  of  the  dorsal  roots  of  the  hooks 
(Fig.  10,  r).  This  part  of  the  rostellum  is  composed  of  minute 
muscle  fibers  which  are  both  circular  and  oblique.  The  rostellum  is 
slightly  broader  below  the  circle  of  hooks  as  it  is  an  oval  shaped 

body. 

The  rostellar  sac  (Fig.  10,  rs)  is  a  deeply  stained  structure  10 
to  I2fi  thick.  It  extends  from  10/x  below  the  hindermost  part  of 
the  rostellum  to  the  anterior  extremity  of  the  scolex,  forming  an 
oval  shaped  sac  or  sheath.  It  is  composed  of  parenchymatous 
tissue  with  large  heavily  stained  oval  or  spindle  shaped  cells  which 
bear  processes.  The  outer  part  of  the  sac  is  composed  of  a  thin 
layer  of  fine  fibers  which  help  to  give  it  a  definite  shape.  At  the 
lower  edges  of  the  sac  the  fil)ers  are  connected  or  associated' to 
some  extent  with  similar  fibers  that  form  the  inner  layer  of  the 
suckers.  The  anterior  region  of  the  rostellar  sac,  which  forms 
the  sheath  for  the  free  head  portions  of  the  rostellum,  is  constructed 
of  an  inner  layer  of  fine  fibers  and  an  outer  layer  of  large  spindle- 
shaped  cells,  the  most  of  which  bear  fibrous  processes  at  one  or 

both  ends. 

The  suckers  are  composed  of  large  spindle-shaped  cells  which 
are  arranged  perpendicular  to  the  edge.     These  are  heavily  stained 


S2  JOHN    E.    GUTBERLET 

and  form  a  compact  layer.  The  inner  boundary  of  the  suckers 
is  composed  of  a  layer  of  fibers  which  are  both  circular  and  oblique. 
Some  of  these  at  the  upper  edges  are  associated  with  similar  fibers 
in  connection  with  the  rostellar  sac. 

The  cyst  is  composed  of  two  cell  layers  with  an  irregular 
cavity  between  them.  The  cells  are  large  and  irregular  in  shape 
with  no  special  arrangement  in  the  layer.  Large  intercellular 
spaces  lie  between  the  cells,  thus  forming  a  loose  network  structure, 
except  at  the  base  of  the  neck.  At  this  point  where  the  neck  is 
attached  to  the  inner  layer  of  the  cyst  the  cells  are  smaller  and 
are  in  a  compact  mass.  There  is  no  definite  boundary  to  the  outer 
part  of  the  inner  layer  as  well  as  to  the  inner  part  of  the  outer 
layer  of  the  cyst.  Few  cells  with  long  connective  processes  extend 
across  the  cavity  from  one  layer  to  the  other.  This  then  forms 
an  irregular  cavity  (Fig.  11  ca)  2  to  20ijl  in  width  between  the  two 
layers  of  the  cyst.  This  is  the  primitive  cavity  of  Grassi  and 
Rovelli  (1889:  373).  The  two  layers  of  the  cyst  are  formed 
apparently  by  a  fold  which  extends  upward  and  inward  from  the 
base  of  the  neck,  forming  the  gastrula  cavity  of  Grassi  and  Rovelli 
(1889:  402,  g)  and  enclosing  the  scolex.  This  cavity  varies  in 
width  from  3  to  10  or  15/i. 

The  bladder,  an  oval  shaped  structure,  is  located  at  one  side 
of  the  cyst  and  is  attached  to  it  at  the  posterior  end  by  a  narrow 
connection  (Fig.  12,  en).  The  posterior  end  of  the  cyst  or  the 
region  caudad  of  the  base  of  the  neck  is  somewhat  drawn  out 
(Fig.  12).  From  this  point  is  given  off  the  attachment  to  the 
bladder  or  tail  portion  of  the  cysticercoid.  The  fact  that  this 
bladder  is  really  a  tail,  even  though  it  possesses  a  cavity,  is  shown 
by  the  presence  of  the  onchospheric  hooks,  which  are  located  at 
the  end  of  the  bladder  opposite  to  that  of  the  attachment  of  the 
cyst   (Fig.  12,  oh). 

The  order  of  arrangement  of  the  onchospheric  hooks  is  indi- 
vidual. In  some  specimens  they  are  situated  at  the  end  of  the 
bladder,  while  in  others  thev  are  at  the  side.  In  some  the  arrang-e- 
ment  is  in  a  group,  while  in  others  they  are  in  pairs.  Some  of 
my  specimens  show  a  pair  of  embryonic  hooks  in  the  layers  of 


MORPHOLOGY   OF    CESTODES    FROM    POULTRY 


33 


the  cyst  between  the  base  of  the  neck  and  the  attachment  of  the 
bladder,  while  the  other  two  pairs  of  hooks  are  located  in  the 
bladder. 

The  cavity  of  the  bladder  is  formed  apparently  by  a  splitting 
or  hollowing  out  of  the  cells  of  the  tail,  because  the  wall  is  con- 
tinuous and  of  the  same  histological  structure.  The  wall  of  the 
bladder  is  constructed  of  two  layers,  an  inner  cell  layer  and  an 
outer  cuticular  layer.  The  outer  cuticular  layer  is  more  or  less 
striated  on  account  of  minute  fibrils  uniting  it  with  the  inner  cell 
layer.  Histologically,  the  structure  of  the  inner  layer  is  con- 
structed of  somewhat  granular  substance  arranged  in  fibers  form- 
ing a  network  which  encloses  clear  spherical  cells  with  large  nuclei 
(Fig.  13).  Outside  of  the  cuticular  layer  is  located  the  peritoneum 
of  the  host  which  lies  upon  the  bladder  and  surrounds  it  as  well 
as  the  cyst. 

C.      COMPARISON    OF   ADULT  AND   CYSTICERCUS 

A  comparative  study  of  the  adult  and  the  cysticercoid  shows 
the  likeness  which  exists  between  them.  The  presence  of  the 
same  number  of  hooks,  having  exactly  the  same  size  and  shape 
as  seen  by  comparing  Figures  8  and  9.  Minute  booklets  of  the 
same  size  are  present  in  both  cysticercoid  and  adult  lining  the  suck- 
ers, the  entire  surface  of  the  scolex  and  a  part  of  the  neck  region. 
Rosseter  (1891:  365)  shows  that  the  hooks  on  the  rostellum  and 
suckers  of  Echlnocotylns  Rosscteri  undergo  no  changes  during 
the  act  of  transition  from  cysticercus  to  adult  stage.  The  rpstellar 
sac  is  of  the  same  general  shape  in  both.  The  head  of  the  rostel- 
lum is  not  expanded  in  the  cysticercoid  as  in  the  adult  because  it 
has  not  functioned  as  yet.  This  corresponds  to  figures  as  shown 
by  Schmidt  (1894.  PI.  VI,  Fig.  A)  of  the  cysticercoid  and  Krabbe 
(1869,  Fl.  VI,  Fig.  114)  of  the  adult  of  Drepanidotmiia  anatma, 
and  by  Grassi' and  Rovelli  (1892,  PI.  IV.  Fig.  7,  8)  of  the  cysti- 
cercoid and  Blanchard  (1891:  16)  of  the  scolex  of  Davainea 
proglottina.  No  measurements  are  given  for  the  rostellum  of 
either  the  cysticercoid  or  the  adult  by  the  above  authors. 

There  is  a  great  deal  of  difiference  in  the  size  of  the  scolex 
between  the   cysticercoid   and   the   adult.     In    my    specimens    the 


34 


JOHN    E.    GUTBERLET 


scolex  of  the  adult  is  between  four  and  five  times  as  large  as  that 
of  the  cysticercoid.  The  scolex  of  the  cysticercoid  has  as  yet 
not  functioned  so  that  the  musculature  of  the  organs  is  not  devel- 
oped as  in  the  adult,  consequently  is  not  nearly  as  massive.  The 
cells  also  are  smaller  than  those  of  the  adult. 

Schmidt  (1894:  10,  44)  shows  that  the  adult  scolex  of  Dre- 
panidotcBnia  anatina  is  about  three  times  as  large  as  that  of  the 
cysticercoid.  He  also  states  that  the  size  of  the  cysticercoid  may 
vary  with  the  size  of  its  host. 

Different  forms  become  modified  in  changing  from  the  inter- 
mediate to  the  adult  hosts  as  shown  by  Schmidt  (1894)  in  Dre- 
panidotcrnia  anatina,  Rosseter  (1891)  in  Echinocotylus  Rosseteri, 
and  Grassi  and  Rovelli  (1892)  in  Davainea  proglottina. 

Onchospheric  hooks  in  the  wall  of  the  tail  are  the  same  size 
(18/a)  and  shape  as  those  of  the  embryos  found  in  the  mature 
proglottids. 

A  consideration  of  these  factors  of  morphological  significance 
which  demonstrate  the  resemblances  between  the  cysticercoid  and 
adult,  indicates  clearly  that  this  cysticercoid  is  the  intermediate 
stage  of  Clioanotcrnia  infundibuliformis. 

OTHER    CHICKEN    CESTODES    IN    THE    UNITED    STATES 

1.     Davainea  tetragona  (Molin  1858)  Blanchard  1891 
Diagnosis:     Length  10  to  250  mm.  by  1  to  2.5  mm.  in  breadth, 
varying  with  state  of  contraction.     Scolex   (Fig.  19)    175  to  215;Lt 
in  diameter,  with  retractile  rostellum  25  to  50/z  in  diameter,  armed 
with  single  row  of  about  100  hooks.     Rostellar  hooks    (Fig.  20) 
6  to  9/i  long  through  longest  axis,  hammer-shaped,  with  long  ven- 
tral root  and  short  dorsal  root,  prong  short  and  recurved.     Suckers 
oval,  60  to  110/1  in  diameter,  armed  with  8  to  10  rows  of  small 
hooks  of  various   sizes.     Acetabular  hooks    (Fig.    21)     range    in 
size     from    4    to    8fi    through    longest    axis,    having    thorn-like 
prong,  short  dorsal  root,  and  longer  flattened  ventral  root,  which 
is  shorter  than  prong.     Neck  long  and  slender,  but  often  as  broad 
as  head.     Segments   trapezoidal  and   imbricate,   edges   of   strobila 
serrate.     Oldest   segments  usually   longer   than  broad,   often  bell- 


MORPHOLOGY   OF    CESTODES    FROM    POULTRY  35 

shaped.  Genital  pores  usually  unilateral,  situated  one  in  each 
segment,  at  or  in  front  of  middle  of  lateral  margin,  frequently 
marked  off  by  papilla.  Male  and  female  canals  pass  on  dorsal 
side  of  nerve  and  excretory  vessels. 

Male  Reproductive  Organs:  Testes  20  to  30  in  median  field 
surrounding  female  organs,  most  of  them  lying  on  aporose  side 
of  latter.  Vas  deferens  situated  in  anterior  third  of  segment, 
beginning  near  median  line,  and  extending  in  much  convoluted 
course  laterally  to  base  of  cirrus  pouch  which  it  enters  and,  after 
a  few  coils  in  basal  portion  of  latter,  passes  into  cirrus.  Cirrus 
pouch  pyriform,  75  to  100/;t  in  length.  Basal  portion  surrounded 
by  prominent  layer  of  longitudinal  muscle  fibers,  neck  with  thick 
layer  of  transverse  fibers.     Cirrus  without  apparent  spines. 

Female  Reproductive  Organs:  Ovary  in  middle  of  segment. 
Yolk  gland  posterior  to  ovary,  irregularly  reniform,  slightly  longer 
in  its  transverse  axis,  about  lOO/x  in  diameter.  Shell  gland  promi- 
nent, 50/i,  in  diameter,  immediately  in  front  of  yolk  gland.  Vagina 
begins  at  genital  pore,  posterior  to  opening  of  cirrus  pouch,  at 
first  very  slender  but  at  distance  of  15  to  25/*  from  genital  pore 
swells  out  into  thick-walled  tube,  functioning  as  seminal  recep- 
tacle. This  extends  transversely  across  segment  and  joins  oviduct 
on  dorsal  side  of  ovary  near  median  line.  Oviduct,  after  being 
joined  in  shell  gland  by  vitelline  duct,  proceeds  forward  and  ends 
on  dorsal  side  of  ovary.  Definite  and  persistent  uterus  not  devel- 
oped. Eggs  pass  from  distal  end  of  oviduct,  become  imbedded 
in  fibrous  and  granular  or  gelatinous  mass  which  fills  up  most  of 
segment.  This  mass  divides  into  50  to  100  portions  to  form  egg 
capsules,  each  surrounded  by  membrane  and  containing  6  to  12 
or  more  eggs.  Egg  is  surrounded  by  three  envelopes, — inner,  close 
to  onchosphere,  often  scarcely  visible ;  middle  layer  or  envelope 
much  folded,  giving  appearance  of  network  between  inner  and 
outer  membranes ;  and  smooth  outer  envelope.  The  onchosphere 
measures  10  to  15/u,  in  diameter;  the  outer  envelope  measures  from 
25  to  50/i  in  diameter. 


One  point  noted  here  that  has  not  been  mentioned  before  by 
other  authors  is  that  the  genital   pores   are   irregularly   alternate. 


36  JOHN    E.    GUTBERLET 

They  are  usually  unilateral.  The  existence  of  this  irregularly 
alternate  occurrence  of  the  genital  pores  may  be  an  anomaly,  but 
it  is  rather  frequent  for  such  a  condition. 

2.  Davainea  echinobothrida  (Megnin  1880)  Blanchard  1891 
Diagnosis :  Length  up  to  250mm ;  width  1  to  4  mm.  Head 
(Fig.  22)  0.25  to  0.45  mm.  in  diameter,  with  retractile  rostellum 
100  to  150/i  in  diameter,  armed  with  crown  of  about  200  hooks 
arranged  in  two  rows.  Suckers  round  or  oval,  90  to  200ju,  in  diam- 
eter, armed  with  8  to  10  rows  of  hooks.  Rostellar  hooks  (Fig. 
23)  similar  to  those  of  Davainea  tctragona,  but  larger,  measuring 
10  to  13)11  in  length.  Acetabular  hooks  (Fig.  24)  likewise  similar 
to  those  of  D.  teiragona,  but  also  larger;  size  variable,  smallest 
being  7  or  S^u,  in  length  and  largest  measuring  from  14  to  16ju,. 
Neck  thicker  and  generally  shorter  than  D.  tetragona,  nearly  equal 
to  width  of  head.  Strobila  resembling  that  of  D.  tetragona,  but 
serrate  border  more  pronounced.  Oldest  segments  in  preserved 
specimens  also  differ  from  those  of  D.  tetragona,  being  less  elon- 
gate and  frequently  marked  by  median  constriction.  Owing  to 
this  constriction  adjacent  borders  of  most  posterior  segments  pull 
apart  in  median  line  and  remain  joined  only  at  sides,  giving  rise 
to  median  series  of  openings  through  posterior  portion  of  strobila. 
Genital  pores  irregularly  alternate,  or  sometimes  almost  entirely 
unilateral,  situated  one  in  each  segment  posterior  to  middle  of 
lateral  margin.  Male  and  female  canals  pass  on  dorsal  side  of 
nerve  and  excretory  vessels. 

Male  Reproductive  Organs:  Testes  20  to  30,  arranged  in 
median  field  surrounding  female  glands  as  in  D.  tetragona.  Vas 
deferens  lies  in  anterior  third  of  segment  much  as  in  D.  tetragona. 
Cirrus  pouch  flask-shaped,  130  to  180ju.  in  length.  Basal  portion 
globular  or  ovoid,  surrounded  by  layer,  about  10;U,  thick,  of  longi- 
tudinal muscle  fibers  inside  of  which  is  a  layer  about  12/ji  thick 
of  transverse  fibers.  Neck  of  pouch  measures  50/u,  to  75ju,  in  length 
by  15  to  20/x  in  diameter,  surrounded  by  layer  of  transverse  fibers 
thickened  at  distal  end  to  form  sphincter.  According  to  Megnin, 
the  cirrus  is  armed  with  minute  spines. 

Female  Reproductive  Organs'.  Female  organs  same  as  in 
Davainea  tetragona,  and  onchospheres    (Fig.  25)   are  also  similar 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  37 

in  Structure  and  size,  14  to  ISju,  in  diameter.  Onchospheric  hooks 
6  to  7/1  long.  Egg  capsules  in  groups  of  6  to  12  or  more,  em- 
bedded in  a  fibi'ous  gelatinous  mass. 

In  the  living  specimens  very  little  difference  can  be  noticed 
except  in  size  of  the  species  D.  tetragona  and  D.  echinohothrida. 
They  are  both  quite  transparent  and  appear  much  alike  in  every 
respect  in  external  appearance,  except  that  D.  tetragona  is  slightly 
more  transparent,  while  the  oldest  segments  of  D.  echinohothrida 
have  very  distinct  median  constrictions  between  them,  appearing 
almost  as  a  series  of  openings. 

The  chief  differences  between  D.  tetragona  and  D.  echinohoth- 
rida are  that  in  the  latter  the  animal  is  larger,  the  hooks  are  more 
numerous  and  larger,  and  the  structure  and  size  of  the  cirrus 
pouches  show  a  very  distinct  difference.  There  is  also  a  difference 
in  the  pathological  effect  of  these  spiny-suckered  forms.  D.  echino- 
hothrida produces  large  nodules  or  ulcers  in  the  intestinal 
wall.  The  scolex  bores  through  the  mucosa  of  the  intestine  and 
in  some  cases  nearly  through  the  muscular  coats.  This  disease 
in  fowls  is  termed  "nodular  t^eniasis",  as  described  by  Moore 
(1895:  1),  and  is  often  mistaken  for  other  diseases. 

3.     Davainea  cesticillus  (Molin  1858)    Blanchard   1891 

Diagnosis:  Length  10  to  125  mm.  Maximum  width  1.5  to 
3  mm.  Head  cylindrical  (Fig.  28),  sometimes  spheriodal,  0.3  to 
0.6  mm.  wide  and  0.2  to  0.4  mm.  long.  Suckers  unarmed,  about 
100/x  in  diameter.  Rostellum  broad  and  flat  or  hemispherical,  0.25 
to  0.35  mm.  wide,  armed  with  a  crown  of  200  to  300  hooks  which 
are  very  unstable  and  easily  lost,  arranged  in  two  ranks.  Hooks 
(Fig.  29)  8  to  12/1  long  with  short  dorsal  root  and  long  ventral 
root.  Neck  very  short.  Anterior  segments  three  to  five  times 
as  broad  as  long;  the  following  increase  in  size  until  they  become 
equal  in  length  and  breadth  and  finally  even  longer  than  broad; 
borders  overlapping.  Genital  pores  irregularly  alternate,  one  in 
each  segment,  somewhat  in  front  of  middle  of  lateral  margin  in 
young  segments  and  nearer  the  middle  in  older  segments.  Vagina 
and  cirrus  pouch  pass  dorsal  of  the  two  excretory  canals  and 
,  nerve. 


38  JOHN    E.    GUTBERLET 

Male  Reproductive  Organs:  Testes  (Fig.  17,  t)  20  to  30  in 
number  in  posterior  portion  of  segment.  Vas  deferens  much 
coiled  before  entering  base  of  cirrus  pouch,  also  coiled  within  latter. 
Cirrus  pouch  ellipsoidal,  120  to  150//,  long  by  55  to  70/t  wide. 
Cirrus  when  protracted  10/t  in  diameter,  armed  with  minute  spines, 
and  with  bulbous  enlargement  20/n  in  diameter  at  its  base,  where 
it  becomes  continuous  with  cirrus  pouch. 

Female  Reproductive  Organs :  Vagina  enlarged  before  reach- 
ing median  line  into  small  seminal  receptacle  (Fig.  17,  sr).  Ovary 
occupies  middle  field  in  front  of  testes.  Yolk  gland  and  shell 
gland  posterior  to  ovary,  ventral  and  dorsal,  respectively,  in  rela- 
tive position.  Uterus  at  first  in  front  of  ovary  as  cord  of  cells ; 
gradually  increasing  in  size,  finally  occupies  most  of  segment  and 
frequently  extends  laterally  beyond  excretory  canals.  In  oldest 
proglottids  it  becomes  divided  into  compartments,  or  capsules, 
each  containing  a  single  egg.  Embryo  (Fig.  30)  36  by  27(1  in 
diameter,  with  very  thin  membrane  closely  adherent  to  surface. 
Embryo  further  enveloped  by  thicker,  smooth  fibrous  membrane, 
oval  in  shape,  45  to  40/a  in  diameter,  with  filament  at  each  pole 
attaching  to  thin  outer  wrinkled  membrane  about  35  by  50/t  in  diam- 
eter :  finally  egg  is  surrounded  by  capsule  composed  of  outer  and 
inner  membrane,  latter  closely  adherent  to  or  fused  with  outer 
egg  membrane ;  and  former  more  or  less  widely  separated  from 
latter  and  connected  with  it  by  number  of  septa. 


One  of  the  principal  points  noted  here  that  is  not  mentioned 
by  other  authors  is  the  size  of  the  rostellar  hooks.  In  my  speci- 
mens they  seem  to  be  somewhat  larger  than  those  described  by 
others.  They  have  been  described  as  being  8  to  lO/x  long,  while 
my  forms  show  many  of  them  to  be  distinctly  I2fi  in  length.  A 
second  point  noted  here  is  the  method  of  the  development  of  the 
uterus.  The  uterus  develops  in  front  of  the  ovary.  It  first  ap- 
pears as  a  solid  cord  of  cells  connected  with  the  united  ducts  of 
the  ovary,  shell  gland,  and  yolk  gland.  The  solid  cord  of  cells 
which  later  gives  rise  to  the  uterus  becomes  hollow  and  appears 
as  a  blind  sac  or  tube.  This  then  grows  in  size,  forming  pockets, 
and  finally  fills  up  the  entire  proglottid. 


MORPHOLOGY   OF   CESTODES    FROM    POULTRY  39 

This  form  is  one  of  the  most  common  chicken  tapeworms 
and  is  the  most  easily  recognized.  It  can  be  identified  by  the  head 
with  its  broad,  flat  rostellum  which  shows  up  very  prominently; 
the  width  of  the  most  anterior  segments  is  usually  equal  to  or 
greater  than  the  width  of  the  head,  and  the  eggs  are  distributed 
in  individual  egg  capsules  in  mature  proglottids. 

4.     Hymenolepis  carioca   (Magalhaes   1898)    Ransom   1902 

Diagnosis:  Length  30  to  80  mm.  Breadth  at  neck  75  to  150/a, 
at  posterior  end  0.5  to  0.7  mm.  Segments  three  to  five  times  or 
more  broader  than  long  throughout  strobila.  Head  (Fig.  26) 
flattened  dorso-ventrally,  140  to  160iLi  long,  150  to  215/x.  wide  and 
100  to  140^  thick.  Suckers  shallow,  70  to  90/x  in  diameter,  un- 
armed. Rostellum  unarmed;  in  retracted  position  25  to  40/x  in 
diameter  and  90  to  100/x  in  length,  with  small  pocket  opening  to 
exterior  in  anterior  position.  Unsegmented  neck  portion  of  strobila 
0.6  to  1.5  mm.  long.  Genital  pores  almost  entirely  unilateral,  a 
single  pore  being  located  in  each  segment  slightly  in  front  of  middle 
of  right-hand  margin. 

Male  Reproductive  Organs:  Testicles  three  in  number,  nor- 
mally two  on  left  and  one  on  right  of  median  line.  On  dorsal 
side  of  inner  end  of  cirrus  pouch  vas  deferens  is  swollen  into 
prominent  seminal  vesicle  (Fig.  18,  sv)  which  may  attain  a  size 
of  70  by  50jLt.  Cirrus  pouch  (Fig.  18,  cp)  in  sexually  mature 
segments  120  to  175^^  long  by  15  to  18/^  in  diameter ;  almost  cylindri- 
cal, slightly  curved  toward  ventral  surface  of  segment;  on  outer 
surface  about  20  longitudinal  muscle  bands,  2  to  3/x  in  thickness, 
very  prominent  in  cross  section;  vas  deferens  enlarged  within 
cirrus  pouch  to  form  small  seminal  reservoir  occupying  proximal 
two-thirds  of  pouch ;  distal  third  of  portion  of  vas  deferens  within 
pouch  very  slender,  about  1/x  in  diameter  and  functions  as  cirrus. 
Genital  cloaca  12  to  36/i  deep. 

Female  Reproductive  Organs:  Opening  of  vagina  in  floor  of 
genital  cloaca,  ventral  and  posterior  to  cirrus  opening.  First  por- 
tion of  vagina  very  narrow,  V  in  diameter.  Small  vaginal  sphinc- 
ter 8  to  10/x  from  vaginal  opening.  On  inner  side  of  sphincter 
vagina   gradually    increases   in    diameter,    and    in    sexually   mature 


40  JOHN    E.    GUTBERLET 

segments  swollen  into  prominent  seminal  receptacle  (Fig.  18,  sr) 
which  extends  forward  to  anterior  border  of  segment  and  inward 
considerable  distance  beyond  proximal  end  of  cirrus  pouch.  Ovary 
faintly  bilobed  or  trilobed  in  posterior  half  of  proglottid.  Yolk 
gland  spherical  or  ovoid,  30  to  40/x  in  diameter,  situated  near  median 
line  of  segment,  posterior  and  dorsal  of  ovary.  Uterus  at  first 
solid  cord  of  cells  extending  transversely  across  segment  along 
anterior  border  of  ovary;  becomes  hollowed  out  and  grows  back- 
ward on  dorsal  side  of  ovary;  in  gravid  segments  occupies  nearly 
entire  segment  and  filled  with  eggs.  Eggs  (Fig.  27)  in  gravid 
uterus  spherical  or  oval,  with  four  thin  membranes,  the  two  middle 
membranes  often  approximate  to  form  thick  layer  which  shows 
somewhat  of  a  cellular  or  coarse  granular  structure.  Diameter  of 
outer  membrane  36  by  36/i  to  75  by  70[i,  of  outer  middle  mem- 
brane 30  by  SOfx  to  65  by  6O/-1,  of  inner  middle  membrane  26  by 
26;u  to  40  by  35/u,,  of  inner  membrane  24  by  16/^  to  29  by  2lfi.  This 
membrane  often  lies  so  close  to  onchosphere  that  it  can  scarcely 
be  distinguished  from  edge  of  embryo.  Onchosphere  is  18  by  14 
to  27  by  19/A  in  diameter;  length  of  embryonal  hooks  10  to  12;U,. 
This  form  is  thread-like  and  usually  occurs  in  great  numbers. 
It  is  very  delicate  and  fragile  and  can  be  recognized  by  that  fact 
alone,  as  it  is  the  most  fragile  of  the  chicken  forms  known. 

SUMMARY 

1.  By  morphological  comparison  of  the  cysticercoids  produced 
experimentally  in  flies  and  adult  of  Choanatocnia  infundibuliformis 
they  are  shown  to  be  identical. 

2.  Morphological  points  noted  are  the  presence  of  minute 
booklets  on  the  suckers  and  entire  surface  of  scolex  in  Choanatcenia 
infundibuliformis.  The  manner  of  development  of  uterus  in  the 
same  species  is  by  means  of  a  blind  tube  which  grows  in  size, 
forming  pockets,  and  later  breaks  up  into  small  compartments. 
In  Davainea  tctragona  the  genital  pores  were  found  to  occur  irreg- 
ularly alternate  in  the  proglottids.  The  hooks  on  the  rostellum 
of  Davainea  cesticillus  were  found  to  vary  in  length  from  8  to 
12ju..  The  uterus  in  development  first  appears  as  a  solid  cord  of 
cells  which  becomes  hollow  and  in  growing  forms  pockets,  fiUing 
the  entire  proglottid. 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  41 

BIBLIOGRAPHY 
Blanchard,  R. 

1891.  Notices  helmiiithologiques.     Sur  les  teniades  a  ventouses  armees. 
Mem.  soc.  zool.  France.,  4:   420-489. 

Davaine,  C. 

1877.  Traite  des  entozoaires  et  des  maladies  vermineuses  de  I'homme  et 
des  animaux  domestiques.    Paris.     Ed.  2,   1003  p. 

Grassi,  B.,  and  Rovelli,  G. 

1888.  Bandwurmer  Entwickelung.  I.  Centralbl.  Bakt.  und  Parasitenk., 
3:  173. 

1889,  Embryologische  Forschungen  an  Cestoden.  Centralbl.  Bakt.  und 
Parasitenk.,  5:  Z70-Z77 ;  401-410. 

1892.  Ricerche  embriologiche  sui  Cestodi.  Atti.  Accad.  Gioenia  Sci. 
Nat.  in  Catania,  4:  1-108. 

Gutberlet,  J.  E. 

1916.     Studies  on  the  Transmission  and  Prevention  of  Cestode  Infection 
in  Chickens.     (In  Press.) 
Hassall,  a. 

1896.  Bibliography  of  Tapeworms  of  Poultiy.  Bull.  Bur.  An.  Ind.,  12: 
81-88. 

Krabbe,  H. 

1869.  Bidrag  til  Kundskat  om  Fuglenes  Baendelorme.  Vid  Selsk.  Skr. 
V.  Roekke.  Nat.  og  Math.,  8:  251-368. 

Magalhaes,  p.  S.  de 

1898.    Notes  d'helminthologie  Bresilienne.    Arch.  Parasit.,  1:  442-451. 

MOLIN,  R. 

1858.  Prospectus  helminthum,  quae  in  prodromo  faunae  helminthologicas 
Venetiae  continentur.  Sitzber.  k.  Akad.  Wiss.  Wien,  math,  naturw. 
kl.,  30:  127-158. 

Moore,  V.  A. 

1895.    A  Nodular  Taeniasis  in  Fowls.    Bur.  An.  Ind.  Cir.  3;  4  pp. 

Mrazek,  Al. 

1907.  Cestoden  Studien.  I.  Cysticercoiden  aus  Lumbriculus  variegatus. 
Zool.  Jahrb.,  Syst.,  24:  591-624. 

PlANA,  G.  p. 

1882.  Di  una  nuova  specie  di  Tenia  del  gallo  domestico  (Taenie  bothri- 
oplitis)  e  di  un  nuova  cisticerco  delle  lumachelle  terrestri  (Cysti- 
cercus  bothrioplitis).    Mem.  Accad.  Sci.  Inst.  Bologna,  2:  387-394. 


42  JOHN    E.    GUTBERLET 

Ransom,  B.  H. 

1900.  A  new  Avian  Cestode-Metroliasthes  lucida.  Trans.  Amer.  Micr. 
Soc,  21 :  213-226. 

1902.  On  Hymenolepis  carioca  (Magalhaes)  and  H.  megalope  (Nitzsch) 
with  Remarks  on  the  Classification  of  the  Group.  Trans.  Amer. 
Micr.  Soc,  23 :  151-172. 

1904,  The  Tapeworms  of  American  Chickens  and  Turkeys.  Ann.  Report 
Bur.  An.  Ind.,  21 :  268-285. 

1904a.  Manson's  Eye-worm  of  Chickens  (Oxyspirura  Mansoni).  Spiny- 
Suckered  Tapeworms  of  Chickens.     Bull.  Bur.  An.  Ind.,  60 ;  12  pp. 

1909.  The  Taenoid  Cestodes  of  North  American  Birds.  Bull.  U.  S.  Nat. 
Mus.,  69:  1-141. 

1911.  A  New  Cestode  from  an  African  Bustard.  Proc.  U.  S.  Nat.  Mus., 
40:  637-647. 

ROSSETER,  T.  B. 

1890.  Cysticercoids  parasitic  in  Cypris  cinerea.  Jour.  Micr.  Nat.  Sci.,  9: 
241-247. 

1891.  Sur  un  cysticercoide  des  Ostracodes,  capable  de  se  developper  dans 
I'intestin  du  canard.    Bull.  soc.  zool.  France,  16:  224-229. 

1892.  On  a  New  Cysticercus  and  a  New  Tapeworm.  Journ.  Queckett 
Micr.  Club,  4:  361-366. 

1897.  On  Experimental  Infection  of  Ducks  with  Cysticercus  coronula 
Mrazek  (Rosseter),  Cysticercus  gracilis  (von  Linstow),  Cysti- 
cercus tenuirostris  (Hamann).  Journ.  Queckett  Micr.  Club,  6: 
397-405. 

Schmidt,  J.  E. 

1894.  Die  Entwicklungsgeschichte  und  der  anatomische  Bau  der  Taenia 
anatina  (Krabbe).    Arch.  Naturg.,  1:  65-112. 

Stiles,  C.  W. 

1896.  Report  upon  the  Present  Knowledge  of  the  Tapeworms  of  Poultry. 
Bull.  Bur.  An.  Ind.,  12;  78  pp. 

TOWEB,  W.  L. 

1900.  The  Nervous  System  of  the  Cestode  Monezia  Expansa.  Zool. 
Jahrb.  Anat.,  13:  359-384. 


MORPHOLOGY    OF    CESTODES    FROM    POULTRY  43 

EXPLANATION  OF  PLATES 

Unless  otherwise  stated  all  drawing  were  made  with  the  aid  of  a  camera 
lucida. 

Abbreviations 

&— bladder  rj— rostellar  sac 

c — cyst  jy— shell  gland 

ca — primitive  cavity  sr — seminal  receptacle 

en — connection  of  bladder  with  cyst     sv — seminal  vesicle 

cp — cirrus  pouch  / — testes 

dex — dorsal  excretory  canal  m — uterus 

ex — excretory  ring  in  scolex  v — vitelline  duct 

0 — ovary  va — vagina 

oh — onchospheric  hooks  vd—\2.%  deferens 

01) — oviduct  vex — ventral  excretory  canal 

r— rostellum  y— yolk  gland 

Plate  V 

CHOANOTAENIA  INFUNDIBULIFORMIS 

Fig.  1.  Scolex  much  contracted.    x40 

Fig.  2.  Scolex  normal  extension.     xl45 

Fig.  3.  Longitudinal  section  of  scolex,  showing  rostellum  and  rostellar  sac. 

x425 

Fig.  4.  Section  of  portion  of  sucker,  showing  booklets.    x425 

Fig.  5.  Section  of  portion  of  wall  of  scolex,  showing  booklets.    x425 

Fig.  6.  Longitudinal  nerve  tract,  showing  nerve  cells  with  processes.     x650 

Plate  VI 

Fig.     7.     A,  B,  C,  D.     Embryos  from  mature  proglottid.     x425 
Fig.    8.     Hooks  from  rostellum  of  adult.    x425 

CYSTICERCUS  OF  CHOANOTAENIA  INFUNDIBULIFORMIS 

Fig.     9.     Hooks  from  rostellum  of  cysticercus.     x42S 

Fig.  10.     Section  through  scolex,  showing  rostellum  with  hooks  and  rostellar 

sac.    x425 
Fig.  11.     Section  through  scolex  and  cyst,   showing  suckers  with  booklets, 

structure  of  cyst  and  primitive  cavity  between  layers  of  cyst.     x425 
Fig.  12.     Reconstruction  of  cysticercus  with  cyst  and  bladder  or  tail,  showing 

scolex  in  cyst  and  onchospheric  hooks  in  bladder.     xl45 
Fig.  13.     Section    of    wall    of    bladder,    showing    histological    structure    and 

peritoneum  of  host.     x42S 


44  JOHN    E.    GUTBERLET 

Plate  VII 

Fig.  14.  Choanotaenia  mfundibuliformis.  Reconstruction  of  mature  pro- 
glottid, showing  reproductive  organs,  excretory  vessels,  and  nerve. 
xl45 

Fig.  15.  C.  infundibuliformis.  Reconstruction  of  cirrus  pouch  showing 
cirrus  and  vas  deferens,  also  part  of  vagina  in  connection  with 
cloaca.    x310 

Fig.  16.  C.  infundibuliformis.  Reconstruction  of  female  reproductive  or- 
gans, showing  part  of  ovary,  yolk  gland,  shell  gland,  oviduct, 
vitelline  duct,  uterus,  and  connection  of  ducts  with  uterus  and 
seminal  receptacle.    x310 

Fig.  17.  Davainea  cesticillus.  Reconstruction  of  mature  proglottid,  showing 
reproductive  organs  and  excretory  vessels.    xl45 

Fig.  18.  Hymenolepis  carioca.  Reconstruction  of  mature  proglottids,  show- 
ing reproductive  organs  from  ventral  view.    xl4S 

Plate  VIII 

Fig.  19.     Scolex  of  Davainea  tetragona.    xl45  •» 

Fig.  20.     Hooks   from  rostellum  of   D.  tetragona.    x425 

Fig.  21.     Hooks  from  suckers  of  D.  tetragona.    x425 

Fig.  22.     Scolex  of  Davainea  echinobothrida.     xl45 

Fig.  23.     Hooks  from  rostellum  of  D.  echinobothrida.    x42S 

Fig.  24.     Hooks   from  suckers  of  D.  echinobothrida.     x425 

Fig.  25.  Embryos  of  D.  echinobothrida,  showing  capsule  and  fibrous  gel- 
atinous mass  in  which  it  is  embedded.    x42S 

Fig.  26.     Scolex  of  Hymenolepis  carioca,  after  Ransom. 

Fig.  27.  A,  B,  C,  D.  Embryos  of  Hymenolepis  carioca,  showing  enveloping 
membranes.     x425 

Fig.  28.  Scolex  of  Davainea  cesticillus.  Free-hand  drawing  of  living  spec- 
imen well  extended,  showing  rostellum. 

Fig.  29.     Hooks  from  rostellum  of  D.  cesticillus.    x425 

Fig.  30.  A,  B,  C,  D.  Embiyos  of  D.  cesticillus,  showing  enveloping  mem- 
branes.   x425 


Plate  V 


^  s 


Plate  VI 


f  ?  7 


29 


1/ 
20 


T 


21 


Contributions  from  the  Zoological  Laboratory  of  the  University  of  Illinois, 
under  the  Direction  of  Henry  B.  Ward,  No.  62. 


\ 


STUDIES  ON  THE  TRANSMISSION  AND  PREVENTION 
OF  CESTODE  INFECTION  IN  CHICKENS 


John  E.  Gutberlet,  Carroll  College,  Waukesha,  Wis. 

Introduction.  The  problem  of  tapeworm  infection  in  chickens 
has  received  but  little  attention  in  the  United  States.  '  In  fact  it 
was  entirely  untouched  until  a  few  years  ago  when  the  subject  was 
opened  by  Stiles  (1896)  and  work  was  begun  by  Eansom  (1900, 
1902,  1904,  1909)  on  poultry  and  other  birds.  At  the  present  time 
less  than  a  dozen  references  constitutes  the  entire  American  lit- 
erature on  the  subject.  Five  species  of  cestodes  are  known  to  in- 
fest chickens  in  various  parts  of  the  United  States. 

No  work  has  been  done  on  the  life  history  of  the  forms 
existing  in  this  country.  However,  studies  have  been  carried 
on  extensively  with  poultry  cestodes  in  various  other  parts  of  the 
world,  though  as  yet  very  little  has  been  finally  determined.  In 
only  one  species  of  chicken  eestode  has  the  life  cycle  been  demon- 
strated experimentally.  That  is  Davainea  proglottina  (Davaine) 
for  which  Grassi  and  Rovelli  (1889 :  372 ;  1892 :  30,  85)  have  shown 
that  the  intermediate  host  is  a  slug  (Limax  cinereus).  This  species 
of  eestode  has  not  as  yet  been  reported  in  this  country. 

Chickens  are  supposed  to  become  infested  with  another  species 
through  eating  snails,  a  third  through  eating  flies,  and  a  fourth 
through  eating  earthworms.  Plana  (1881-1882)  found  in  a  snail 
(Helix)  two  cysticereoids  which  agree  closely  with  the  head  of 
Davainea  tetragona  (Molin).  No  experiments  were  performed  to 
demonstrate  that  the  cysticercoid  was  the  larval  stage  of  that 
species  and  the  only  evidence  of  their  connection  is  the  similarity 
in  form.  Grassi  and  Rovelli  (1892:  33,  87)  found  in  flies  cysticer- 
eoids which  closely  resembled  Choanotaenia  mfundibuliformis  and 
base  their  conclusion  of  identity  on  the  structural  similarity. 

Grassi  and  Rovelli  (1889  :  372  ;  1892 :  29)  found  in  earthworms 
(Allolhophora  foetida)  cysticereoids  which  they  associated  with  the 
scolex  of  Dicranotaenia  sphenoides,  a  chicken  eestode  not  reported 
in  this  country.  Here  again  the  only  evidence  for  regarding  it  to 
be  the  larval  stage  of  this  species  is  a  general  structural  likeness. 
In  no  one  of  these  three  forms  was  the  life  cycle  demonstrated  ex- 
perimentally. Such  comparisons  are  not  proof  that  the  cysticer- 
eoids are  intermediate  stages  of  definite  species,  but  only  give  a 
clue  as  to  the  probable  life  cycle. 


CESTODE  INFECTION   IN   CHICKENS  219 

In  other  kinds  of  poultry  more  has  been  done  on  the  life  his- 
tories of  their  eestodes.  The  life  cycles  of  five  species  of  duck 
cestodes  have  been  demonstrated  through  experiment.  Schmidt 
(1894)  proved  that  Drepanidotaenia  anatina  (Krabbe)  has  its  in- 
termediate stage  in  a  fresh-water  crustacean  (Cypris  ovata).  He 
fed  large  quantities  of  tapeworm  eggs  to  the  crustaceans  and  found 
that  the  larvae  developed  in  two  weeks  during  the  summer.  Ros- 
seter  (1891,  1892)  has  shown  that  a  second  duck  cestode,  Echino- 
cotylus  Rosseteri  (Blanchard),  has  its  intermediate  stage  in  an- 
other small  fresh-water  crustacean  (Cypris  cinereus).  He  fed  large 
numbers  of  the  crustaceans  to  ducks  which  upon  examination  later 
yielded  a  large  crop  of  tapeworms  of  the  species  named. 

Jlosseter  (1897)  also  demonstrated  experimentally  the  life  his- 
tories of  three  other  species  of  duck  cestodes.  He  had  discovered 
some  cysticerci  in  crustaceans  which  he  compared  with  the  adult 
<vorms  occurring  in  ducks  and  found  that  they  agreed  closely.  He 
produced  Dicranotaenia  coronula  in  a  duck  by  feeding  it  Cypris 
cinerea.  Drepanidotaenia  gracilis  was  introduced  into  the  ducks 
through  Cypris  cinerea  and  Cypris  viriens.  Drepanidotaenia 
temiirostris  was  likewise  raised  by  feeding  Cyclops  agilis. 

As  in  other  cases  the  question  of  control  of  infection  in  chickens 
depends  to  a  great  extent  upon  the  life  history  of  the  parasites. 
Little  can  be  done  to  wipe  out  the  disease  until  more  is  known  of 
its  source.  Certain  methods  may  be  employed  to  check  it,  but  as 
yet  it  has  been  impossible  to  prove  the  exact  source  of  infection. 
Usually  it  is  easiest  to  control  such  forms  during  the  developmental 
stages. 

This  paper  is  the  result  of  some  investigations  carried  on  to 
find  out  the  life  history  of  certain  chicken  tapeworms.  Numerous 
experiments  were  tried  on  various  insects  and  many  observations 
made  on  the  habits  of  the  birds  in  the  endeavor  to  ascertain  where 
the  cause  of  the  infection  was  located.  The  habits  of  the  birds  are 
probably  the  chief  factors  to  be  dealt  with  in  experiments  of  this 
kind.  Certain  insects  that  are  common  about  the  habitats  of  the 
birds  are  readily  eaten.  They  are  hence  more  likely  to  be  inter- 
mediate hosts  than  those  which  are  rare  in  these  localities.  Such 
factors  have  been  taken  into  consideration  and  through  experiment 
it  has  been  shown  that  one  cestode,  Choanotaenia  infundibuli- 
formis,  has  its  intermediate  stage  in  the  common  house-fly. 

The  most  of  the  material  was  collected,  and  the  experimental 


220  JOHN  E.  GUTBERLET 


work  was  done  on  a  farm  at  Hardy,  Nebraska.  A  large  amount  of 
material  was  also  collected  at  the  poultry  farm  at  the  University 
of  Illinois. 

Thanks  are  due  to  Professor  D.  0.  Barto,  of  the  University 
of  Illinois,  for  giving  me  the  privilege  of  collecting  material  at 
the  poultry  farm.  For  other  assistance  I  am  indebted  to  my  father 
and  mother,  William  and  Flora  Gutberlet,  for  their  untiring  efforts 
to  make  this  work  a  success  by  taking  records  and  making  collec- 
tions of  material  at  times  of  the  year  when  they  would  not  otherwise 
have  been  taken. 

I  wish  to  express  my  appreciation  to  Dr.  Henry  B.  Ward,  at 
whose  suggestion  this  work  was  first  taken  up,  for  his  helpful  sug- 
gestions and  criticisms  during  the  preparation  of  this  paper. 

Methods  op  Technic.  In  making  collections  of  tapeworms 
the  intestine  of  the  bird  was  slit  open  under  water  and  the  con- 
tents removed  by  shaking  gently.  The  worms  are  usually  attached 
to  the  wall  and  can  be  easily  seen  and  removed  with  the  aid  of  a 
pair  of  needles.  Those  that  are  not  attached  sink  to  the  bottom  of 
the  dish. 

In  removing  the  worms  from  the  intestine  it  was  found  best 
to  transfer  them  directly  to  fresh  water.  A  weak  saline  solution^ 
was  demonstrated  to  be  harmful  as  the  worms  die  in  it  in  a  very 
short  time.  Tower  (1900:  362)  found  saline  solution  harmful  to 
cattle  cestodes  (Monezia).  In  fresh  water,  the  worms  soon  become 
well  extended  and  remain  alive  and  normal  for  twelve  to  fifteen 
hours,  or  even  longer.  The  worms  are  best  killed  in  a  corrosive- 
acetic  solution  and  preserved  in  70%  alcohol  and  glycerine.  For 
study  of  structure  and  accurate  diagnosis  of  species  the  worms 
were  cut  in  sections  from  5  to  10  microns  in  thickness,  stained  in 
Delafield's  or  Ehrlich's  acid  haematoxylin  and  destained  in  acid 
alcohol. 

In  order  to  use  house-flies  for  experimental  purposes  one  has 
to  work  out  first,  methods  of  keeping  them  alive.  The  flies  used 
were  kept  for  experiment  in  small  cages.  They  demanded  a  great 
deal  of  attention  because  the  slightest  disturbance  of  conditions 
was  harmful.  They  were  fed  most  satisfactorily  on  blood,  liver, 
and  spleen.  It  was  found  that  a  fly  could  not  live  long  without- 
a  constant  supply  of  water  in  the  cages.  The  cages  had  also  to  be 
placed  in  the  sun  for  a  few  minutes  each  morning,  and  then  kept 
in  the  shade  for  the  rest  of  the   day,  but  not  in  a  cool  place. 


CESTODE   INFECTION   IN   CHICKENS 


221 


At  the  conclusion  of  the  experiment  the  flies  were  killed,  fixed  in 
corrosive-acetic  solution  and  preserved  in  70%  alcohol.  The  chitin 
covering  of  the  body  of  the  flies  was  punctured  to  allow  these 
fluids  to  penetrate  properly. 

Large  bottles  proved  very  satisfactory  as  cages  for  beetles 
during  the  experiments.  The"  bottles  were  fitted  with  glass  or 
metal  stoppers  provided  with  pores  for  the  passage  of  air.  Leaves 
and  a  small  amount  of  soil  were  placed  in  the  bottom  of  the  bottle. 
The  beetles  were  killed  and  preserved  like  the  flies,  but  before 
sectioning  the  chitin  covering  was  removed  by  dissection. 

Amount  op  Infection.  The  flock  of  chickens  upon  which 
these  studies  were  carried  on  was  so  heavily  infected  with  the  tape- 
worm disease  during  certain  seasons  that  it  was  rather  unusual  to 
find  a  bird  that  did  not  harbor  at  least  a  few  of  the  parasites.  The 
investigations  extended  over  a  period  of  two  summers.  Close  ob- 
servations were  made  during  those  seasons,  and  also  at  several 
other  times  during  the  year,  to  secure  a  record  of  the  amount  of 
infection  during  other  seasons  than  the  summer  months. 

The  first  summer  (1912)  about  fifty  chickens  were  examined 
for  parasites.  Eight  of  these  were  adults  and  in  no  ease  was  there 
any  nifection.  Ten  young  birds  from  six  weeks  to  two  months  old 
were  examined  in  June,  but  none  of  them  were  infected.  The  first  in- 
fection of  tapeworms  for  that  year  was  detected  on  July  25.  Between 
that  date  and  September  9,  thirty-two  young  birds  were  examined 
and  every  one  showed  some  infection.  In  some  it  was  slight,  while 
'in  others  it  was  very  heavy.  During  this  same  period,  between 
July  25  and  September  9,  some  adult  birds  were  examined  but 
yielded  no  parasites. 

During  the  summer  of  1913  forty  birds  were  examined  between 
August  10  and  September  18,  with  some  infection  in  every  bird. 
A  few  of  these  were  adult  birds  which  had  only  a  few  parasites. 
The  young  birds  were  more  heavily  infected,  although  the  number 
of  parasites  varied  with  different  birds.  In  one  bird  which  was  ex- 
amined at  the  age  of  seven  weeks,  twenty-five  tapeworms  were 
found.  Between  June  17  and  August  1,  eight  birds  were  examined 
and  cestodes  were  present  in  every  bird  with  the  exception  of  one 
adult  killed  on  June  24. 

I  have  records  of  infection  in  the  flock  for  January  1  and 
April  27,  1913,  and  for  November  20,  December  2,  and  December 
26,  1913.  There  are  five  species  of  worms  infesting  the  chickens 
in  this  place  and  further  details  are  given  in  the  table. 


222 


JOHN  E.  GUTBERLET 


Between  June  20  and  August  1,  1913,  examinations  were  made 
of  about  fifteen  birds  at  Urbana,  111.  Some  of  these  were  from 
the  poultry  farm  at  the  University  of  Illinois  and  others  were  from 
private  yards  of  residents  in  this  vicinity;  in  only  one  bird  was 
there  any  trace  of  an  infection.  In  that  case  there  were  a  few 
fragments  of  worms  which  were  in  such  a  state  of  disintegration 
that  they  could  not  be  preserved  or  determined.  No  further  ex- 
aminations for  parasites  were  made  in  this  locality  until  December 
2,  1913,  when  it  was  discovered  that  the  chickens  at  the  poultry 
farm  at  the  University  of  Illinois  were  badly  infested.  Several 
were  examined  and  found  to  harbor  Davainea  echinohothrida, 
Davainea  cesticillus,  and  Hymcnolepsis  carioca. 

A  general  examination  was  made  of  the  living  birds  at  the 
poultry  farm  and  it  was  discovered  that  symptoms  of  cestode  dis- 
ease were  manifested  by  the  great  majority  of  the  chickens,  al- 
though the  infection  was  apparently  not  heavy  except  in  a  small 
percentage  of  the  flock. 

In  making  examinations  upon  dead  birds  infested  with  Dava- 
inea echinohothrida  it  was  found  that  large  nodules  were  formed 
in  the  intestinal  wall  which  is  a  characteristic  pathologic  effect 
of  this  particular  spiny-suckered  form.  Davainea  cesticillus  seems 
to  be  almost  universally  present  as  there  was  hardly  an  infested 
bird  examined  in  Nebraska  or  Illinois  that  did  not  harbor  some  of 
this  species. 

The  following  table  shows  the  amount  of  infection  and  the 
number  of  worms  occurring  in  each  bird  examined,  both  in  Ne- 
braska and  Illinois : 


DATE 

LOCALITY 

Age  of 
host 

No.  of 
D.  cesti- 
cillus 

No.  of 

D.  tetra- 

gona 

No.  of 
D.  echino- 
bothrida 

No.  of 

Hymeno. 

carioca 

No.  of  Ch 

infundibu 

liformis 

July 

1 1 

25, 

1912 
29 

Hardy,  Neb. 

4  mo. 
4  mo. 

5 
2 

8 

August 

2 

7 

5  mo. 
5  mo. 

3 
3 

15 

20 

2 

8 

5  mo. 

10 

25 

10 

9 

5  mo. 

10 

35 

10 

5  mo. 

5 

15 

5 

2 

11 

5  mo 

7 

10 

3 

12 

5  mo. 

10 

10 

30 

3 

15 

5  mo. 

5 

4 

1 

20 

Adult 

20 

5  mo. 

3 

15 

3 

1 

21 

5  mo. 

15 

10 

21 

5  mo. 

3 

2 

2 

22 

5  mo. 

3 

8 

3 

CESTODE  INFECTION   IN   CHICKENS 


223 


No.  of 

No.  of 

No.  of 

No.  of 

No.  of  Ch. 

DATE 

LOCALITY 

Age  of 

D.  cest- 

D.  tetra- 

D.  echlno- 

Hymeno. 

Infundibu- 

Host 

cillus 

gona 

bothrida 

carioca 

liformis 

( < 

23 

t  ( 

5  mo. 

3 

3 

2 

2 

i_( 

25 

( t 

5  mo. 

5 

10 

10 

ti 

27 

I  i 

5  mo 

6 

4 

5 

1 1 

29 

ct 

Adult 

September 

2 

<  t 

Adult 

( ( 

4 

t  ( 

6  mo. 

5 

20 

10 

It 

6 

(  c 

6  mo. 

15 

10 

6 

It 

7 

Hardy 

6  mo. 

10 

14 

5 

1 1 

8 

( < 

6  mo. 

4 

10 

2 

ft 

9 

1 1 

6  mo. 

10 

10 

5 

January  2,  1913 

1 1 

Adult 

3 

5 

March 

23 

i  c 

Adult 

( ( 

25 

i  I 

Adult 

i  t 

30 

1 1 

Adult 

April 

27 

1 1 

Adult 

10 

June 

17 

it 

Adult 

<  I 

24 

t  < 

Adult 

1 1 

21 

Urbana 

2  yr. 

tt 

23 

I  i 

2  yr. 

1 1 

28 

i  I 

2  yr. 

July  7,  17,  20 

24 

Hardy 

4  mo. 

infection,  but  recor 

ds  incomplete. 

August 

10 

I  i 

4  mo. 

5 

10 

15 

1 1 

11 

I  c 

4  mo. 

3 

( e 

15 

i  I 

4  mo. 

3 

i  t 

16 

( t 

5  mo. 

15 

30 

20 

10 

tt 

17 

It 

5  mo. 

25 

30 

5 

8 

tt 

19 

I  ( 

5  mo. 

6 

15 

5 

tt 

20 

1 1 

5  mo. 

6 

6 

tt 

26 

I  c 

5  mo. 

15 

2 

tt 

26 

i  I 

5  mo. 

20 

25 

8 

10 

t  ( 

27 

1 1 

5  mo. 

1 

5 

4 

1 1 

28 

1 1 

5  mo. 

2 

10 

10 

5 

It 

28 

i  ( 

5  mo. 

10 

10 

6 

t  i 

30 

Hardy 

5  mo. 

8 

18 

3 

September 

2 

<  i. 

Adult 

3 

12 

4 

( ( 

4 

<  i 

5  mo. 

20 

20 

( t 

6 

i  c 

7  wks. 

17 

8 

tt 

10 

t  ( 

5  mo. 

8 

10 

5 

ti 

10 

1 1, 

5  mo. 

15 

25 

4 

tt 

11 

I  e 

5  mo. 

10 

15 

4 

tt 

12 

1 1 

5  mo. 

25 

35 

15 

1 1 

13 

1 1 

5  mo. 

8 

10 

4 

8 

It 

15 

t  i 

6  mo. 

10 

10 

5 

1 1 

16 

I  I 

6  mo. 

3 

30 

tt 

17 

1 1 

6  mo. 

20 

10 

1 1 

20 

i  i 

8  mo. 

35 

6 

10 

December 

2 

Urbana 

4  mo. 

20 

30 

<  < 

5 

<  < 

4  mo. 

3 

( t 

6 

<  ( 

Adult 

5 

5 

tt 

9 

( ( 

6  mo. 

6 

6 

1 1 

24 

Knoxville,    111     Adult 

17 

00 

00 

GO 

1 

f  t 

26 

Hardy 

Adult 

1 

11 

3 

January  1,  1914 

( I 

Adult 

3 

2 

5 

224  JOHN  E.  GUTBERLET 


Symptoms  and  Effects  of  Tapeworm  Infection.  A  great 
deal  has  been  written  on  the  symptoms  of  this  disease  by  various 
authors,  but  in  every  case  they  were  unable  to  reach  any  definite 
conclusions  on  the  subject.  In  my  own  study,  which  was  exten- 
sive, I  reached  the  following  definite  conclusions:  The  symptoms, 
while  not  really  individual,  vary  to  some  extent  with  the  different 
birds,  with  the  age  of  the  birds,  and  with  the  degree  of  infection. 
Some  birds  are  affected  by  the  disease  much  more  than  others  and 
show  sj^mptoms  and  effects  much  more  readily.  Some  birds  that 
show  no  symptoms  and  appear  in  good  health  are  heavily  infested 
with  the  worms,  while  others  showing  severe  effects  and  manifesting 
all  the  symptoms  are  not  nearly  as  heavily  infested.  The  age  of  the 
host  is  a  factor  of  much  importance  for  indicating  the  presence  of 
an  infection  with  the  species  I  studied.  Young,  growing  birds  are 
affected  much  more  than  adults  and  show  the  symptoms  more  dis- 
tinctly. Even  a  comparatively  slight  infection  can  be  detected  in 
a  young  bird  a  few  weeks  of  age,  while  a  heavy  infection  is  very 
marked.  Most  adults  manifest  no  external  symptoms  as  far  as 
appearance  is  concerned  unless  they  are  heavily  infested.  The  de- 
gree of  infection  is  another  factor  which  is  of  importance  in 
making  a  diagnosis  for  cestodes.  Birds  that  harbor  only  a  few 
worms  show  conditions  which  are  quite  different  from  those  that 
possess  a  large  number.  Therefore  the  symptoms  are  rather  var- 
iable. 

Stiles  (1896:  13)  mentions  some  general  principles  for  diag- 
nosis, and  Zurn  (1882:  17)  gives  more  fully  some  of  the  symptoms 
that  may  be  taken  as  indications  of  the  disease  in  the  birds. 

In  general,  one  may  say  that  a  light  infection  can  hardly  be 
noticed  and  is  apparently  in  no  way  harmful  to  the  fowl.  In 
cases  suffering  from  a  moderate  to  a  heavy  infection  the  conditions 
were  found  to  be  quite  different.  In  the  first  place,  birds  that  are 
moderately  infested  are  apparently  always  hungry,  having  in- 
deed ravenous  appetites  and  seeming  never  to  be  able  to  get  enough 
to  eat.  Secondly,  they  manifest  a  great  desire  for  water,  increas- 
ing in  cases  where  the  infection  is  heavy.  Moreover,  infected  birds 
are  greedy  and  it  seems  as  if  their  hunger  had  caused  them  to  lose 
control  of  themselves  whenever  there  is  a  chance  to  obtain  any  food. 
Such  birds  are  also  restless,  always  moving  about  as  if  searching 
for  something.  This  in  part  probably  accounts  for  the  fact  that 
the  fowls  are  poor  in  flesh  and  more  or  less  in  an  emaciated  con- 


CESTODE  INFECTION   IN   CHICKENS  225 


dition.  They  are  never  at  ease  on  account  of  their  restless  atti- 
tude which  is  apparently  due  to  nervousness.  Normal  exercise 
alone  does  not  depress  the  condition  of  the  bird,  but  rather  the 
constant  restlessness  and  uneasiness  which  is  manifested  by  those 
that  are  infested. 

The  heavily  infested  chickens  become  emaciated  and  lose  their 
color,  the  feathers  become  ruffled,  and  the  plumage  is  not  glossy  as 
in  the  fowls  that  are  free  from  the  disease.  Growing  birds  that 
were  heavily  infested,  were  found  usually  to  be  slender  and 
quite  poor  in  flesh,  the  head  very  thin  and  the  comb  pale.  In 
cases  of  heavy  infection  the  growing  birds  isolate  themselves  to 
some  extent  and  often  allow  the  wings  to  droop  and  hang  at  the 
sides.  The  sick  birds,  even  though  they  isolate  themselves,  still 
manifest  a  great  desire  for  food  and  water. 

A  slight  infection  is  hardly  to  be  detected  in  the  droppings, 
but  when  it  is  heavy  there  is  developed  an  irritation  or  inflamma- 
tion of  the  intestinal  epithelium,  a  kind  of  catarrh  which  results  in 
a  diarrhea,  varying  with  the  degree  of  infection.  This  irritation 
of  the  intestinal  epithelium  by  the  worms  causes  an  abundant  flow 
of  mucus  into  the  intestine.  The  mucous  secretion  is  at  first  a 
clear,  transparent  semi-liquid,  and  sometimes  slightly  whiH^ish. 
Worms  which  are  slightly  transparent  are  difficult  to  see,  as  they 
are  imbedded  in  the  mucus.  Later  the  mucus  takes  on  a  brownish 
color  which  is  due  in  part  to  slight  hemorrhages  of  the  epithelium 
caused  by  the  irritation  of  the  worms.  This  color  of  the  mucus  is 
retained  until  it  is  passed  out  with  the  feces  so  that  the  droppings 
of  an  infested  bird  have  always  a  characteristic  yellowish-brown 
color.  This  factor  of  coloration  in  the  droppings  is  one  that  can 
nearly  always  be  depended  upon  as  a  criterion  of  infection. 

When  the  infection  is  heavy  a  gas  is  formed  in  the  intestine 
which  is  noticeable  in  the  droppings  in  the  form  of  bubbles.  These 
bubbles  are  present  when  the  feces  are  first  passed  and  remain  in 
the  semi-liquid  droppings  for  some  time.  This  is  very  character- 
istic in  cases  of  heavy  infection  but  is  not  noticeable  at  other  times 
except  in  cases  of  extreme  diarrhea,  and  then  the  gaseous  forma- 
tion is  comparatively  slight.  In  a  flock  that  is  heavily  infested 
nearly  every  dropping  detected  about  roosting  or  resting  places 
shows  the  characteristic  yellowish-brown  color  with  a  large  number 
of  small  gas  bubbles  enclosed.  Tlie  infested  birds  pass  droppings 
often,  though  in  small  quantities. 


226  JOHN  E.  GUTBERLET 


Segments  of  the  worms  can  usually  be  found  when  there  is 
a  moderately  he&Yj  infection,  and  eggs  can  nearly  always  be  dem- 
onstrated by  the  aid  of  a  microscope,  but  the  latter  method  is  not 
practical  under  all  circumstances.  When  the  above  methods  fail 
to  show  any  signs  of  infection  and  an  absolute  diagnosis  is  desired 
it  may  be  well  to  take  a  few  of  the  birds  that  show  some  of  the  symp- 
toms, kill  them  and  make  an  examination  of  the  contents  of  the  in- 
testine between  the  gizzard  and  caecum.  Any  infection  which 
cannot  be  detected  by  the  above  methods  is  so  slight  that  it  is  not 
harmful  to  the  birds  in  any  way,  or  is  so  recent  that  the  cestodes 
are  too  small  to  be  seen. 

The  best  criteria  for  diagnosis  are  the  emaciated  condition  of 
the  birds,  the  great  desire  for  food  and  water,  and  the  marked  diar- 
rhea with  the  characteristic  yellowish-brown  color  of  the  droppings ; 
furthermore  in  cases  of  heavy  infection  segments  of  worms  can 
usually  be  detected,  though  there  is  some  degree  of  uncertainty  in 
making  gross  examinations  for  the  proglottids  in  the  feces.  The 
excretions  from  the  kidneys  are  white  in  color  and  at  times  have 
somewhat  the  appearance  of  the  tapeworm  proglottides.  This  may 
at  times  be  misleading  to  one  who  is  inexperienced  with  this  method 
of  examination.  The  excretion  from  the  kidneys  can  be  readily 
distinguished  from  proglottids  by  placing  the  droppings  in  water 
and  breaking  up  the  mass.  Proglottids  have  a  definite  shape  and 
are  firm,  while  the  excretions  break  up  into  fine  granules,  or  shreds 
which  are  easily  disintegrated  by  shaking. 

Some  of  the  above  symptoms  for  cestode  infection  are  iden- 
tical with  those  for  nematodes :  the  emaciated,  unthrifty  condition, 
the  ruffled,  dull  appearance  of  the  feathers,  and  the  more  or  less 
restless  attitude  of  the  bird.  The  feces,  however,  look  quite  differ- 
ent and  often  blood  is  passed  with  the  droppings  in  cases  of 
nematode  infection.  The  nematodes  produce  hemorrhages  in  the 
intestine  by  boring  into  the  epithelium. 

Tapeworm  infection  is  harmful  according  to  the  degree  of  in- 
fection. A  slight  infection  does  practically  no  harm  to  the  bird, 
but  when  there  is  a  heavy  infection  the  condition  is  more  serious. 
The  intestinal  inflammation  or  catarrh  is  quite  a  serious  matter 
and  in  many  cases  may  prove  fatal.  It  brings  on  a  more  or. less 
anaemic  condition  and  the  bird 's  general  health  is  run  down.  Such 
a  condition  is  suitable  for  the  coming  in  of  other  diseases,  since 
the  fowl  is  unable  to  ward  them  off  because  of  its  weakened  state 


CESTODE  INFECTION   IN   CHICKENS 


227 


of  health.  Through  these  means  the  tapeworms  are  most  harmful, 
as  their  effect  works  more  or  less  indirectly  with  other  diseases. 

I  have  found  instances  where  the  worms  were  so  numerous 
that  they  would  form  such  a  large  compact  mass  in  the  intestine 
as  to  interfere  with  passage.  These  masses  imbedded  in  a  great 
quantity  of  mucus  become  lodged  at  the  junction  of  the  small  and 
large  intestines  with  the  caeca. 

One  species,  Davainea  echinohothrida,  produces  nodules  or 
ulcers  in  the  intestinal  wall  which  are  often  mistaken  for  other 
diseases.  This  has  a  more  serious  effect  upon  the  chickens  than 
some  of  the  other  species  as  it  has  more  of  a  direct  pathological 

effect. 

Chickens  infested  with  any  of  the  species  of  common  tape- 
worms devoured  great  quantities  of  food,  but  upon  examination  the 
intestines  were  usually  found  empty.  It  seems  as  if  the  food 
material  after  reaching  the  intestine  rushes  through  rapidly  on 
account  of  the  large  amount  of  mucus  and  the  marked  diarrhea. 
This  does  not  allow  the  bird  to  obtain  as  much  nourishment  as  it 
would  otherwise.  The  cestodes  of  course  absorb  their  nourishment 
from  the  chyme  in  the  intestine.  Furthermore,  the  excretions  from 
the  worms  may  also  have  some  effect  upon  the  general  health  of 
the  bird,  as  some  are  without  doubt  resorbed  into  the  system  from 
the  intestine. 

More  practical  proof  must  be  obtained  by  experimental  study 
on  the  various  effects  and  symptoms  of  infection  in  chickens  before 
much  can  be  definitely  said  on  the  subject.  As  yet  there  is  but 
little  known  in  regard  to  definite  symptoms  and  effects  except  in  a 
general  way. 

Methods  of  Control.  The  subject  of  the  control  and  treat- 
ment of  tapeworm  disease  in  chickens  has  not  been  studied  exten- 
sively. There  is  need  of  more  experimental  data  before  much  can 
be  said  concerning  it.  Several  remedies,  however,  have  been  tried 
with  some  degree  of  success,  although  they  do  not  seem  practical 
when  large  numbers  of  birds  are  to  be  treated. 

A  practice  general  among  poultry  raisers  is  to  isolate  a  sick 
bird  and  leave  it  to  cure  itself,  or  to  kill  it.  Most  poultry  men  do 
not  take  the  trouble  to  treat  a  sick  bird  nor  do  they  even  try  to 
find  out  the  cause  of  its  ailment,  but  simply  say  that  it  has  "gone 
light."  Such  an  expression  covers  a  multitude  of  diseases  pre- 
valent among  poultry.     Birds  that  are  heavily  infested  with  worms 


228  JOHN  E.  GUTBERLET 


isolate  themselves  and  become  emaciated.     They  are  also  said  to 
have  ''gone  light." 

As  the  first  prerequisite  for  carrying  on  any  sort  of  treat- 
ment for  worm  diseases,  the  infected  birds  must  be  isolated  from 
the  rest  of  the  flock  so  that  the  latter  can  be  kept  free  from  con- 
tamination. The  droppings  from  the  sick  birds  must  be  cared 
for  or  destroyed  in  some  way  so  that  the  embryos  of  the  worms  are 
killed  and  insects  prevented  from  feeding  on  them.  In  general 
for  any  flock,  preventive  measures  should  be  taken  against  infection 
of  all  kinds  by  keeping  the  surroundings  clean  and  sanitary;  all 
droppings  around  roosts  should  be  collected  often  or  subjected  to 
such  treatment  as  will  render  them  harmless  or  inaccessible  to  in- 
sects. "Wherever  an  infection  is  present,  even  if  only  slight,  such 
preventive  measures  should  be  taken  to  eliminate  all  possibility  of 
its  further  increase.  One  of  the  best  is  to  collect  the  droppings'* 
about  the  coop  daily  and  place  them  into  vats  or  cans  that  are  in- 
accessible to  insects  or  worms ;  they  are  then  treated  with  lime 
or  some  substance  which  destroys  the  embryos.  Lime  or  ashes 
should  be  scattered  over  the  droppings  around  the  roosts  and  the 
resting  places  of  the  birds.  This  destroys  the  embryos  and  keeps 
insects  from  feeding  upon  the  droppings.  Furthermore,  if  the 
droppings  are  covered  with  lime  and  collected  often  it  will  prevent 
insects  from  breeding  in  them.  House-flies  especially,  lay  eggs  in 
chicken  manure  if  the  droppings  are  not  treated  with  lime. 

Other  features  in  the  habitat  of  the  birds  should  be  kept 
sanitary ;  such  are  the  feeding  places  and  drinking  vessels.  Water- 
ing troughs  should  be  so  placed  that  the  birds  cannot  get  their  feet 
into  them,  as  they  may  carry  in  eggs  or  embryos  of  other  para- 
sitic worms  (nematodes)  which  will  reach  the  birds  again  through 
the  water  if  the  latter  is  allowed  to  stand  in  a  filthy  condition. 

The  location  of  poultry  yards  should  be  changed  from  time  to 
time  if  possible,  because  if  the  same  grounds  are  used  from  year 
to  year  some  of  the  insects  that  may  be  the  intermediate  hosts  of 
the  tapeworms  may  become  numerous  and  thus  increase  the  possi- 
bility of  infection.  Embryos  of  parasites  or  germs  of  certain  dis- 
eases remain  on  the  premises  from  year  to  year,  and  if  the  yards  are 
changed,  more  healthful  conditions  are  produced  for  the  birds. 

In  addition  to  destroying  the  eggs  and  embryos  of  parasites 
in  the  droppings,  it  is  fully  as  important  to  destroy  the  adult  in- 
sects and  their  breeding  places.     The  life  history  of  only  one  spe- 


CESTODE  INFECTION   IN   CHICKENS 


229 


cies  of  tapeworm  has  been  worked  out  in  the  United  States,  as  is 
discussed  elsewhere  in  this  paper.  This  species  is  known  to  have 
its  intermediate  stage  in  the  house-fly.  House-flies  breed  commonly 
in  bird  or  horse  manure,  or  any  decaying  vegetable  matter.  The 
destruction  of  all  such  breeding  places  is  a  difficult  matter  and 
little  can  be  done  along  that  line  or  with  the  destruction  of  adult 
flies.  However,  fly  traps*  can  be  placed  over  the  windows  of  the 
chicken  coop  and  many  flies  caught  and  killed. 

According  to  Stiles  (1896:18),  the  principal  remedies  that 
have  been  used  for  the  removal  of  tapeworms  from  poultry  are 
such  drugs  as  extract  of  male  fern,  turpentine,  powdered  kamala, 
areca  nut,  pomegranate  root  bark,  pumpkin  seeds,  and  sulphate  of 
copper.  These  have  been  experimented  with  to  a  certain  extent 
and  have  been  found  to  be  satisfactory  in  some  instances. 

The  experiments  with  these  remedies  have  been  worked  out 
on  individual  birds.  Each  bird  must  be  treated  individually. 
While  such  methods  of  treatment  are  thorough,  they  are  not  prac- 
tical for  a  poultry  raiser  who  has  an  infection  in  a  flock  of  several 
hundred  birds.  It  would  require  handling  each  bird  separately 
two  or  three  times,  and  demand  a  considerable  amount  of  time ;  too 
much  to  be  practicable  on  account  of  the  expense  involved. 

I  tried  experiments  on  a  number  of  birds  to  see  whether  a 
more  practical  method  could  be  found.  It  had  been  observed 
previously  that  hogs  infected  with  worms  could  be  freed  from  them 
by  feeding  the  ashes  from  corncobs.  The  ashes  contain  a  large 
amount  of  sodium  and  potassium  carbonate.  Lye  is  made  from  ashes 
and  of  course  contains  similar  substances,  together  with  sodium  hy- 
droxide. 

The  following  experiment  worked  very  successfully :  Fifteen 
birds  which  showed  symptoms  of  tapeworm  infection  were  placed 
in  a  cage  which  was  insect-proof  and  were  given  the  following 
treatment;  A  gallon  of  a  mixture  of  wheat  and  oats,  to  which 
was  added  a  small  tablespoonful  of  concentrated  lye,  was  cooked 
slowly  for  about  two  hours  and  allowed  to  cool.  The  birds  were 
fasted  for  about  fifteen  hours  and  were  then  given  as  much  of  the 
mixture  as  they  would  eat,  with  plenty  of  water.  Twelve  hours 
later  one  of  the  birds  was  killed  and  an  examination  of  the  small 
intestine  was  made.     It  was  found  that  nearly  all  of  the  worms  in 

*Such  as  described  by  F.  C.  Bishopp,  Farmers '  Bulletin  No.  540,  1913. 


230  JOHN  E.  GUTBERLET 


the  intestine  were  loose,  the  scolices  being  detached  from  the  wall, 
and  were  also  apparently  dead.  The  rest  of  the  birds  were  given 
a  second  dose  twenty-four  hours  after  the  first.  Many  worms  had 
passed  with  the  droppings  in  from  twenty-four  to  twenty-six  hours 
after  the  first  feeding.  Most  of  the  worms  in  these  droppings 
were  dead,  but  in  all  probability  the  embryos  were  still  alive  in  the 
mature  proglottids.  Twelve  hours  after  the  second  dose  was  given 
another  bird  was  killed  and  it  was  found  that  only  a  few  worms  were 
left  and  all  of  these  were  detached  and  dead.  The  intestine  was 
filled  with  a  peculiar  gray  colored,  slimy  substance  composed  mainly 
of  mucus.  Many  entire  worms  and  fragments  were  passed  with 
the  droppings  during  the  period  of  the  feeding.  The  lye  acted^to 
some  extent  as  a  purgative. 

The  birds  were  given  normal  diet  again,  and  in  a  few  days 
they  showed  no  symptoms  of  infection.  Eight  days  after  the  sec- 
ond dose  two  more  birds  were  killed  and  examinations  made.  One 
possessed  a  small  fragment  of  a  tapeworm  and  the  other  was  en- 
tirely free. 

The  effects  of  such  treatment  upon  the  flock  as  a  whole  were 
shown  later.  While  I  was  carrying  on  other  investigations  with 
chicken  cestodes  my  father  noticed  that  the  birds  were  very  heav- 
ily infested  with  worms.  In  an  endeavor  to  free  the  birds  of  the 
worms  and  to  improve  their  general  condition  he  fed  them  a  mix- 
ture of  cooked  grain  and  lye  on  July  15,  unknown  to  me.  As  a 
result  the  entire  flock  of  nearly  four  hundred  birds  was  practically 
frejed  from  the  worms  by  a  single  application  of  the  remedy.  The 
cestodes  were  so  thoroughly  removed  that  there  were  not  enough  left 
to  allow  me  to  go  on  with  my  investigations  and  my  observations  on 
the  worms  were  not  taken  up  again  until  August  10,  when  the  birds 
had  become  infested  again  and  the  parasites  had  grown  to  such  size 
as  to  enable  the  continuance  of  my  work. 

This  remedy  is  a  very  simple  one  and  is  practical.  It  has 
been  known  to  many  poultry  raisers  for  some  time,  but  they  have 
neglected  to  use  it,  mainly  on  account  of  the  fact  that  heretofore 
no  definite  evidence  has  ever  been  presented  concerning  its  actual 
working  possibilities.  It  may  not,  and  in  all  probability  will  not, 
remove  all  the  worms,  but  it  does  remove  most  of  them  so  that  they 
are  not  serious  and  can  be  controlled  in  the  flock  as  a  whole. 

In  a  large  flock  the  birds  can  be  housed  for  the  length  of 
time  required  for  the  fast,  then  fed  on  the  cooked  grain  and  kept 


CESTODE  INFECTION   IN   CHICKENS 


231 


in  the  house  until  after  the  effects  of  the  second  dose  have  passed 
off.  During  the  time  that  they  are  confined  the  droppings  should 
be  collected  often  and  lots  of  lime  used  about  the  coop  and  over 
the  droppings  to  keep  away  the  insects.  In  a  flock  the  treatment 
would  have  to  be  repeated  from  time  to  time  whenever  the  birds 
became  infected  again.  Further  experimental  evidence  must  be  ob- 
tained before  much  can  be  said  in  regard  to  details  of  this  method 
of  treatment,  especially  as  to  the  amount  of  the  alkali  to  be  used. 
A  large  amount  would  be  harmful  to  the  intestinal  mucosa,  while  a 
small  amount  would  have  little  if  any  effect  upon  the  parasites. 

Feeding  Experiments  for  Infection.  Chickens  in  the  vicin- 
ity of  Hardy,  Nebraska,  were  heavily  infested  with  tapeworms,  and 
young  birds  were  found  to  be  more  heavily  infested  than  the  adults. 
This  led  to  investigations  concerning  the  reason  for  the  difference 
in  the  infestation  of  the  adult  and  young  birds  when  they  were  to- 
gether in  the  same  environment  and  fed  on  the  same  diet. 

The  summer  of  1913  was  very  dry  in  the  locality  which  was  a 
factor  in  keeping  the  numerous  varieties  of  insects  down  to  a  min- 
imum, because  the  drought  interfered  with  their  breeding.  Upon 
observation  it  was  found  that  only  two  kinds  of  insects  were  present 
in  any  abundance  about  the  haunts  of  the  birds.  Those  were  the 
ground  beetle  Tenehrio  and  flies.  The  stable  fly,  Stomoxys  calci- 
trans,  which  usually  breeds  in  wet,  decaying  straw,  was  very  scarce 
because  its  breeding  places  had  dried  up.  The  house  flies  were 
very  abundant  everywhere. 

The  reason  why  the  adults  should  be  only  slightly  infested 
with  parasites,  while  the  young  and  growing  birds  harbored  so  many, 
was  then  the  subject  for  observation.  The  birds  were  watched  in 
their  haunts  and  their  habits  studied.  It  was  soon  noticed  that 
the  young  birds,  when  in  their  resting  places  in  the  shade  of  a  tree 
or  a  building,  were  busy  the  whole  time  pursuing  flies  and  very 
often  caught  their  prey,  while  the  adults  paid  little  or  no  attention 
to  the  flies.  This  led  to  the  conclusion  that  flies  might  have  some- 
thing to  do  with  the  transmission  of  the  worms  to  the  birds. 

With  a  view  to  testing  this  hypothesis,  experiments  were  car- 
ried on  with  the  worms  that  were  most  common  in  the  birds. 
These  species  were  Davainea  cesticillus,  Davainea  tetragona,  and 
Choanotae7iia  infundibuliformis. 

Segments  of  these  worms  were  teased  apart  so  that  the  eggs  or 
embryos  were  set  free  in  a  drop  of  water,  and  this  was  fed  to  flies 


232  JOHN  E.  GUTBERLET 


of  the  species:    Musca  domestica,  Stomoxys  calcitrans,  and  Calli- 
phora  vomitaria. 

Only  a  few  Calliphora  could  be  obtained  and  these  did  not 
live  long  under  experimental  conditions.  This  species  of  fly  does 
not  frequent  places  where  it  would  be  likely  to  become  the  inter- 
mediate host  of  any  of  the  chicken  cestodes,  as  it  always  remains  in 
cool,  damp,  and  usually  dark  places,  unless  it  can  find  carrion. 
However,  on  cool,  dark,  damp  days  it  does  appear  in  chicken  yards, 
but  its  occurrence  there  is  not  frequent.  Some  Stomoxys  were  used, 
but  in  no  case  did  they  live  long  in  captivity.  -* 

Musca  domestica  lived  much  longer  than  either  of  the  others, 
even  though  it  was  difficult  to  keep  them  alive  for  a  long  period. 
After  a  great  deal  of  experimentation  it  was  found  that  they  could 
be  kept  alive  in  a  cage  for  twelve  or  thirteen  days,  and  in  one  ex- 
treme case  some  were  kept  alive  for  twenty-one  days.  The  flies  in 
captivity  were  fed  on  blood,  liver  and  spleen.  These  were  found  to 
be  the  best  foods. 

The  oldest  proglottids  on  the  worm  were  usually  taken  for 
feeding  to  flies,  and  also  some  of  the  free  segments  in  the  intestine 
were  used.  The  use  of  the  oldest  proglottids  proved  to  be  an  error 
in  the  case  of  Choanotaenia  infundibtdiformis,  because  it  was  found 
later  that  in  this  species  the  oldest  segments  separate  from  the 
worm  before  they  are  entirely  mature,  but  proglottids  that  have  been 
free  in  the  intestine  for  some  time  may  be  mature.  The  use  of 
proglottids  that  were  not  entirely  mature  for  feeding  flies  was  an 
error  in  my  experiments  which  may  account  for  so  few  infections. 

Davainea  cesticillus.  In  a  series  of  experiments  107  flies  of 
the  species  Musca  domestica  were  fed  on  the  eggs  from  proglottids 
of  Davainea  cesticillus.  Some  were  killed  and  preserved  each  day 
from  the  beginning  of  the  experiment  until  the  tenth  day,  when  the 
remaining  flies  died,  except  in  one  case  four  were  kept  alive  for 
twenty-one  days.  These  were  all  sectioned  with  the  exception  of 
five,  which  were  dissected.  No  stages  of  the  cestodes  were  found  in 
any  of  the  flies  when  examined. 

During  the  experiment  microscopic  examinations  were  made  of 
a  great  number  of  the  droppings  of  the  flies  and  no  eggs  or  embryos 
of  the  worms  could  be  found  in  any  case.  It  is  certain  that  the 
flies  got  some  of  the  eggs  because  they  were  numerous  in  the  mater- 
ial that  was  fed  to  them.  The  flies  would  lap  up  all  the  water  in 
which  the  eggs  floated  and  would  then  suck  on  the  fragments  of 


CESTODE   INFECTION   IN   CHICKENS 


233 


proglottids.  Ill  several  instances  when  the  flies  were  hungry  it 
was  observed  that  they  would  take  small  fragments  of  the  proglot- 
tids between  the  labella  of  the  labium  and  actually  devour  them. 
Since  the  eggs  are  microscopic  in  size  it  is  practically  certain  that 
the  flies  got  some  of  them. 

Several  Calliphora  were  fed  on  eggs  from  this  species,  but 
these  flies  lived  for  only  two  or  three  days. 

Proglottids  of  this  tapeworm  were  fed  to  a  number  of  beetles 
of  the  species  Tenehrio  melitor.  The  beetles  ate  the  segments 
readily.  Some  were  killed  at  the  end  of  one  week,  others  at  two 
weeks,  and  the  rest  at  three  weeks.  These  beetles  were  sectioned, 
but  showed  no  developmental  stage  of  cestodes. 

Davainea  tetragona.  In  experiments  on  this  species  59  flies 
in  all  were  used.  Some  of  these  were  killed  and  preserved  after 
from  two  to  twelve  days.  The  proglottids  were  broken  up  and 
the  eggs  set  free  in  a  drop  of  water.  The  flies  lapped  up  the  water 
with  the  eggs  and  afterwards  sucked  all  of  the  moisture  from  the 
^fragments  of  the  proglottids.  Therefore,  it  is  very  probable  that 
the  flies  got  some  of  the  eggs.  Microscopic  examinations  of  the 
droppings  of  the  flies  showed  no  signs  of  eggs. 

Material  would  pass  through  the  flies  in  a  few  hours  as  was 
demonstrated  by  feeding  them  on  blood.  When  the  flies  gorged 
themselves  with  blood  they  passed  red  droppings  in  from  eight  to 
ten  or  twelve  hours.  This  indicated  the  length  of  time  that  it  took 
material  to  pass  through  the  alimentary  canal.  In  this  way  the 
approximate  time  to  make  fecal  examinations  for  the  eggs  was  de- 
termined. However,  examinations  were  made  of  the  droppings 
after  five  or  six  hours  as  well  as  later  and  at  regular  intervals  of 
two  or  three  hours. 

The  flies  were  fed  on  eggs  once  or  twice  each  day  for  three  days. 
When  they  were  fed  once  a  day  that  was  done  in  the  morning,  and 
when  fed  twice  they  were  given  one  dose  in  the  morning  and  the 
other  at  noon.  On  three  occasions  some  flies  were  fed  in  the  even- 
ing and  fecal  examinations  were  made  the  next  morning  and  con- 
tinued at  intervals  of  two  or  three  hours. 

The  flies  were  all  sectioned  and  examined,  but  showed  no  stages 
of  the  cestodes  in  any  instance. 

Some  Calliphora  were  fed  upon  the  eggs  of  this  species,  but 
they  did  not  live  more  than  two  or  three  days.  Some  beetles,  Ten- 
ehrio, were  fed  on  proglottids,  but  upon  examination  they  showed 
nothing. 


234  JOHN  E.  GUTBERLET 


Choanotaenia  infundibuliformis.  Eggs  of  this  species  were 
fed  to  88  flies  of  the  species  Musca  domestica.  Besides  these  some 
Stomoxys  calcitrans  were  also  fed,  but  these  did  not  live  long  in 
captivity.  The  individuals  of  Musca  domestica  used  in  these  ex- 
periments lived  from  two  to  seventeen  days.  Two  flies  lived  for 
twelve  days  and  four  for  seventeen  days.  The  proglottids  were 
broken  up  and  fed  to  the  flies  in  the  same  manner  as  in  the  other 
species  mentioned.  All  of  these  flies  were  sectioned  and  examined. 
One  fly  preserved  at  the  end  of  twelve  days  showed  five  cysticerci. 
These  cysticerci  agree  very  closely  with  the  structure  of  the  adult 
of  this  species,  and  the  hooks  are  identical.  This  cysticercus  is  de- 
scribed in  detail  in  another  paper. 

Grassi  and  Rovelli  (1892:  33)  found  cysticerci  in  flies  which 
they  compared  with  this  species.  They  found  that  there  was  a  close 
agreement  in  structure  between  the  cysticerci  they  discovered  and 
the  adult  of  Choanotaenia  infundihidiformis.  They  therefore  in- 
ferred that  the  larva  they  had  was  the  intermediate  stage  of  this 
species,  but  did  not  demonstrate  experimentally  its  connection  with 
the  adult  tapeworm. 

During  the  process  of  my  experiments  I  had  hoped  to  be  able 
to  feed  some  chicks  on  flies  that  had  been  previously  fed  on  tape- 
worm eggs,  but  as  it  was  so  difficult  to  keep  the  flies  alive  under  ex- 
perimental conditions  such  an  experiment  could  not  be  carried  out. 
However,  another  feeding  experiment  was  tried  with  the  following 
results :  Six  chicks  were  taken  from  the  nest  as  soon  as  they  were 
hatched  and  placed  in  a  cage  where  they  could  get  no  insects  and 
great  care  was  taken  during  feeding  so  that  no  flies  could  enter. 
Flies  (Musca  domestica)  were  caught  around  the  chicken  roosts  and 
fed  to  three  of  the  chicks.  The  other  three  birds  were  used  as  a 
control  and  were  given  no  flies.  Fifty  flies  were  fed  to  each  of 
the  three  chicks.  Three  weeks  after  feeding,  the  chicks  were  killed 
and  examined  with  the  result  that  two  were  found  to  be  infested 
with  Choanotaenia  infundibidiformis.  One  bird  possessed  six 
worms.  These  were  of  the  same  length,  being  35  mm.  long,  and  each 
one  contained  103  proglottids.  The  other  bird  had  one  worm  of 
the  same  species,  but  it  was  a  little  longer,  43  mm.  and  having  118 
proglottids.  This  bird  was  fed  on  the  flies  three  days  before  the 
one  sheltering  the  six  worms.  The  three  birds  which  were  used  as 
a  check  on  the  experiments  contained  no  worms  when  killed  and  ex- 
amined. 


CESTODE  INFECTION  IN   CHICKENS  235 


These  six  birds  were  kept  together  in  a  cage  and  were  fed  on 
corn  meal  and  bread  crumbs.  The  three  birds  that  were  fed  flies 
were  caught  and  the  insects  were  given  to  them  from  the  hand. 

A  number  of  stable  flies  (Stomoxijs  calcitrans)-weve  used  in 
the  experiments  with  this  species  of  worm,  but  they  would  not  live 
under  experimental  conditions  for  any  length  of  time.  They  would 
usually  die  within  24  to  36  hours,  except  in  one  case  when  six  lived 
for  five  days.     They  were  sectioned,  but  nothing  could  be  found. 

On  numerous  occasions  I  have  observed  maggots  in  the  drop- 
pings beneath  the  chicken  roosts.  Now,  since  house-flies  are  in  the 
habit  of  breeding  in  such  places,  it  seemed  possible  that  infection 
might  take  place  in  the  maggot  stage  of  the  flies.  Experiments 
were  then  tried  with  the  maggots  of  Musca  domestica  and  Stomoxys 
calcitrans.  Thirty  Musca  domestica  maggots  were  fed  on  segments 
of  three  species  of  cestodes,  Davainea  cesticillus,  Davainea  tetra- 
gona,  and  Choanotaenia  infimdihuliformis.  The  maggots  devel- 
oped puparia  in  a  day  or  two.  Some  were  sectioned  in  the  pupa 
stage.  The  rest  developed  into  adults  and  were  sectioned,  a  few 
were  dissected,  but  only  negative  results  were  obtained.  Fifty 
maggots  of  Stomoxys  calcitrans  were  fed  on  proglottids  of  the  same 
three  species  of  tapeworms.  The  maggots  went  into  the  pupal  stage 
within  two  or  three  days.  Some  were  sectioned  in  the  pupal  stage. 
Most  of  them  developed  into  adults  and  were  sectioned  while  a  few 
were  dissected.  No  positive  results  were  obtained  from  either  pupal 
or  adult  stages. 

From  the  foregoing  it  seems  probable  that  flies  are  not  the 
intermediate  hosts  for  Davainea  cesticillus  and  Davainea  tetragona, 
as  the  experiments  that  I  have  carried  on  with  them  are  extensive 
enough  to  appear  conclusive.  However,  the  small  number  of  vari- 
eties of  insects  present  in  the  locality  seems  to  throw  the  burden 
upon  the  flies,  since  they  were  so  abundant  and  observations  show 
that  they  are  taken  and  eaten  by  the  chickens  that  are  most  heavily 
infested.  The  adult  birds  eat  all  other  insects  that  are  easy  to 
catch,  but  since  the  flies  are  more  difficult  to  take  as  prey  they 
leave  them  alone.  If  the  infection  is  direct,  the  adults  would  have 
fully  as  much  chance  as  the  young  birds  because  they  get  food  and 
water  together  and  have  the  same  environment. 

In  the  case  of  Choanotaenia  infundibuliformis  it  seems  to  be 
clear  that  the  house-fly  is  the  intermediate  host.  Grassi  and 
Rovelli  hold  that  it  is  the  intermediate  host  on  a  purely  structural 
basis.    My  experiments  show  that  it  is  certainly  an  intermediate 


236  JOHN  E.  GUTBERLET 


host  in  some  cases.  Furtherwore,  feeding  chicks  on  flies  that  were 
taken  from  about  the  chicken  roosts  and  raising  the  cestodes  make 
it  probable  that  the  house-flies  are  the  intermediate  hosts  of  this 
one  species. 

The  reason  why  more  flies  were  not  infected  by  feeding  on 
the  eggs  of  this  species  was  determined  to  be  peculiar  conditions  in 
the  maturing  of  the  proglottid.  At  the  time  when  the  experiments 
were  being  carried  on  it  was  not  known  that  the  oldest  proglottids 
separated  from  the  worm  before  they  are  entirely  mature.  In  the 
experiments  the  oldest  proglottids  on  the  worm  were  usually  taken 
for  feeding,  though  in  some  cases  the  free  segments  in  the  intestine 
were  used.  Since  the  flies  were  fed  on  eggs  that  were  not  entire- 
ly mature  the  embryos  were  digested.  The  free  proglottids  remain 
in  the  intestine  of  the  bird  for  some  time  and  in  all  probability 
mature  there.  Some  such  free  proglottids  were  examined  and  found 
to  contain  mature  embryos. 

SUMMARY 

1.  The  results  of  these  experiments  show  that  the  intermediate 
(cysticercoid)  stage  of  Choanotaenia  infundihuliformis  occurs  in  the 
common  house-fly  Musca  domestica.  The  results  were  obtained  by 
feeding  flies  on  eggs  of  the  tapeworm  and  raising  cysticercoids  in 
a  fly ;  also  by  feeding  chicks  on  flies  and  raising  the  worms  in  the 
birds.  By  morphological  comparison  of  the  cysticercoid  and  adult 
they  are  shown  to  be  identical.  Results  from  experiments  by  feed- 
ing flies  on  eggs  from  Davainea  cesticillus  and  Davainea  tetragona 
were  negative. 

2.  The  habits  of  the  birds  are  important  factors  to  be  con- 
sidered in  experimental  work  for  life  history  studies.  Certain  in- 
sects are  found  in  great  numbers  around  chicken  houses  and 
yards  and  are  readily  eaten  by  the  birds.  Flies  are  known  to  con- 
tain the  larval  stage  of  one  species  of  cestode,  and  some  other 
species  of  insects  are  to  be  considered  as  probable  intermediate 
hosts  for  other  species  of  cestodes. 

3.  The  symptoms  and  effects  of  the  infection  from  tapeworms 
vary  with  individual  birds,  age  of  birds,  and  the  degree  of  infec- 
tion. Birds  infested  with  worms  display  an  emaciated,  unthrifty 
condition,  an  unnatural  desire  for  food  and  water,  and  a  marked 
diarrhea  with  droppings  of  a  characteristic  yellowish-brown  color. 

4.  The  control  of  tapeworm  disease  in  chickens  is  in  an  unset- 
tled condition.  Little  can  be  done  until  more  is  known  concerning 
life  histories  of  worms.    Preventive  measures  are  urged  rather 


CESTODE  INFECTION   IN   CHICKENS  237 

than  curative  measures.  Droppings  should  be  cared  for  and  treated 
with  appropriate  substances  in  order  to  prevent  insects  from  feed- 
ing on  them  or  developing  in  them.  Experiments  by  giving  lye 
with  food  to  infested  chickens  showed  satisfactory  results  in  re- 
moving tapeworms, 

5.  The  flocks  of  chickens  that  were  studied  showed  at  times  a 
very  heavy  infection  and  nearly  every  bird  examined  harbored  one 
or  more  species  of  worms.  Five  species  were  found  in  the  chickens 
at  Hardy,  Nebraska,  and  three  in  the  birds  at  the  poultry  farm  at 
the  University  of  Illinois.  The  species  found  in  Nebraska  are 
Davainea  cesticillus  (Molin),  Davainea  tetragona  (Molin),  Davainea 
echinohothrida  (Megnin),  Hymenolepis  carioca  (Magalhaes),  and 
Choanotaejiia  infundibuliformis  (Goeze).  At  the  poultry  farm  of 
the  University  the  species  Davainea  cesticillus  (Molin),  Davainea 
echinohothrida  (Megnin)  and  Hymenolepis  carioca  (Magalhaes) 
were  found. 

6.  A  full  description  of  the  structure  of  these  parasites  has 
been  published  in  the  Transactions  of  The  American  Microscopical 
Society,  Vol.  35,  p.  23-44,  PI.  5-8. 


BIBLIOGEAPHY 

Grassi,  B.  B.  and  Eovelli,  G.     1889.     Embryologische  Forschungen  an  Ces- 
toden.     Centralhl.  f.  BaM.  und  Parasitenl:,  5:370-377,  401-410. 
1892.     Kicerche  embriologiche  sui   Cestodi.     Atti  Accad.   Gioenia  di  Set. 
Nat.  in  Catania,  4:1-108. 
PiANA,  G.  P.     1882.    Di  una  nuova  specie  di  Tenia  del  gallo  domestico  (Taenia 
bothrioplitis)    e   di  un  nuova  cisticerco   delle  luniachelle   terrestri    (Cysti- 
cercus  bothrioplitis).     Mem.  Accad.  Set.  Inst.  Bologna,  2:387-394. 
Eansom,  B.  H.     1900.     A  new  Avian  Cestode— Metroliasthes    lucida.     Trans. 
Amer.  Micr.  Soc,  21:213-226. 

1902.     On  Hymenolepis  carioca   (Magalhaes)   and  H.  megalops   (Nitzsch) 
with   Remarks   on   the   Classification   of   the   Group.    Trans.  Amer.  Micr. 
Soc.,  23:  151-172. 
1904.     The  Tapeworms  of  American  Chickens  and  Turkeys.     Bur.  An.  Ind. 

Ann.  Bpt.,  21 :  268-285. 

1909.     The  Taenoid  Cestodes  of  North  American  Birds.    Bull.  U.  S.  Nat 

Mus.,  69:  1-141. 

EossETER,  T.  B.     1891.     Sur  un  Cysticercoide  des  Ostracodes,  capable  de  se  de- 

velopper  dans  I'intestin  du  Canard.     Bull.  Soc.  Zool,  France,  16:  224-229. 

1892.     On   a  New   Cysticercus   and   a   New   Tapeworm.     Journ.    Queckett 

Micr.  Club,  4 :  361-366.. 
1897.     On   Experimental    Infection    of    Ducks   with    Cysticercus    coronula 
Mrazek  (Rosseter),  Cysticercus  gracilis  (von  Linstow),  Cysticercus  tenui- 
rostris  (Hamann).     Journ.  Queekett  Micr.  Club,  6:  397-405. 
Schmidt,  J.  E.     1894.     Die  Entwicklungsgeschiehte  und  der  anatomische  Bau 

der' Taenia  anatina  (Krabbe).     Areh.  f.  Naturg.,  1894,  1:  65-112. 
Stiles,  C.  W.     1896.     Report  upon  the  Present  Knowledge  of  the  Tapeworms 

of  Poultry.     Bur.  An.  Ind.  Bull.  No.  12;  78  pp. 
Tower,  W.  L.     1900.     The  Nervous  System  of  the  Cestode  Monezia  Expansa. 

Zool.  Jahrb.,  13 :  359-384. 
ZtJRN,  F.  A.     1882,    Die  Krankheiteu  des  Hausgefliigels.     237  pp.,   76  figs. 
Weimar. 


I 


i 


VITA 

John  Earl  Gutberlet  was  born  in  Nebraska  in  March,  1887. 
He  received  his  preparatory  training  in  the  high  school  at  Hardy, 
Nebraska,  and  at  Bethany  Academy,  Lindsborg,  Kansas.  He 
entered  Bethany  College,  Lindsborg,  Kansas,  in  1905  and  received 
the  A.  B.  degree  from  that  institution  in  1909.  During  the  year 
1909-10  he  was  a  graduate  student  and  an  assistant  in  the 
department  of  Zoology  at  the  University  of  Colorado.  He 
attended  the  summer  session  of  the  University  of  Colorado 
Biological  Station  in  1910.  In  the  fall  of  1910  he  went  to  the 
University  of  Illinois  where  he  accepted  a  position  as  assistant 
in  the  department  of  Zoology  which  he  held  until  1913.  He 
received  the  A.  M.  degree  from  the  University  of  Illinois  in  1911. 
During  the  sunmier  of  1911  he  was  engaged  in  study  at  the  Marine 
Biological  Station,  La  JoUa,  California.  He  was  elected  to  mem- 
bership in  the  Illinois  Chapter  of  Sigma  Xi  in  1912.  He  attended 
the  summer  session  of  the  University  of  Illinois  in  1913.  During 
the  year  1913-14  he  held  a  fellowship  in  Zoology  at  the  Univer- 
sity of  Illinois  where  he  completed  his  work  for  the  degree  of 
Doctor  of  Philosophy  in  June,  1914. 


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